Effects of fragmentation on artificial nest predation in a tropical forest in Kenya

Nest predation has been established as a leading source of reproductive failure in songbirds. Numerous studies, conducted primarily in the Northern Hemisphere, have linked high rates of nest predation with fragmentation and habitat edges. To date, virtually no such studies have been conducted in Tropical Africa. Here we investigate the effects of disturbance level and edge on depredation of artificial avian nests in six fragments of Kakamega Forest, Kenya. We found that nest losses were higher at the edge than in the interior as often reported in previous studies. Surprisingly, nest predation was highest in the least disturbed fragments. One possible explanation for these findings is that top predators are missing from Kakamega Forest, allowing nest predator populations to increase in all fragments, while in the highly disturbed fragments, domestic dogs might be surrogates for top predators. Alternatively, human activity may itself lead to a decline in mesopredator populations.     

Effects of habitat fragmentation on pollen flow and genetic diversity of the endangered tropical tree Swietenia humilis (Meliaceae)

Fragmentation of tropical forest represents a major threat to some tree populations by reducing local population size and gene flow from other populations. Both processes can decrease outcrossing rates and genetic variation in remnant stands. Despite these risks, some tree species have pollen vectors that mitigate these negative consequences for fragmented populations. In this paper, we assess both pollen flow and diversity of pollen sources in continuous forest and isolated stands of Swietenia humilis, a tropical tree species pollinated by small insects. Using seven nuclear microsatellite markers, we test the hypothesis that genetic diversity and the number of pollen donors are lower in remnant populations. Results show that allelic richness of seeds is lower in isolated populations (6.1 vs. 8.3 alleles per locus), even though adult populations do not show this difference.Pollen pool structure is greater in isolated patches (Φ_Iso = 0.26) than in continuous forest (Φ_For = 0.14), which yields estimates of the average effective number of pollen donors (N_ep) of 1.9 and 3.6 respectively. In addition, estimates of number of sires per mother indicate that isolated trees have half the number of pollen sources (4.98) than trees in the forest (9.8). Although extensive pollen movement (>2000 m) was recorded on both habitat conditions, indicating that fragmented patches are not isolated from pollen-mediated gene flow, this extensive pollen flow among trees in fragmented landscapes may not serve to counteract deleterious reproductive and genetic consequences of habitat fragmentation.     

Effects of forest fragmentation, anthropogenic edges and fruit colour on the consumption of ornithocoric fruits

We investigated effects of fruit colour (red, black or white), habitat (anthropogenic edges and forest interior) and fragment size on the removal of artificial fruits in semideciduous forests in south-east Brazil. Eight forest fragments ranging from 251 to 36,000 ha were used. We used artificial fruits, which were placed on shrubs between 1 and 2 m in height and checked after 48 and 96 h for peck marks in the fruits. All three variables affected the probability of consumption of our fruit models. Red and black fruits were statistically more pecked than the white fruits. The probability of fruit consumption was lower in the interior than at the edge and less in small than in large fragments. However, the decrease fruit consumption in small compared with large fragments was more accentuated for red and black fruits than for white fruits. Our results show that habitat reduction and edges affect the chances of a fruit being eaten by birds, which may ultimately affect plant fitness in forest fragments.     

Nest predation in a fragmented Afrotropical forest: evidence from natural and artificial nests

Nest predation accounts for a substantial share of nest failure and low reproductive success in most tropical songbirds. Normally, forest fragmentation leads to an increase in nest predation pressure due to reduced cover, fewer (and poorer) nest sites and predator influxes from the surrounding habitats. To test this hypothesis, we studied natural nesting behaviour and nest success of the white-starred robin (Pogonocichla stellata) in seven Afrotropical forest fragments differing in size and level of habitat disturbance. Based on data from 12 nests, we estimate that 29% of all natural nests initiated by the robins survive to produce fledglings across all fragments. We also conducted an experiment using artificial (plasticine) model-eggs to reveal potential predators and compare relative predation rates amongst fragments. This experiment revealed that small mammals might be the major predators on robin nests at the egg-stage. In addition, it showed that the highest incidences of nest disturbance during this stage were in the most heavily disturbed fragment. This was presumably attributable to an influx of mammalian predators from the surrounding habitats as forest degradation created suitable habitats for them. Such an infiltration was recently reported in this study site. Both nest placement and microhabitat did not significantly affect depredation levels in our experiment. This suggests that depredation was predominantly incidental (i.e., predators mainly encountered nests fortuitously while foraging for other food items), where the likelihood of encountering a nest largely depended upon the prevalence of the principal potential predators - the small mammals.     

The effects of forest fragmentation on euglossine bee communities (Hymenoptera: Apidae: Euglossini)

Despite scientific and media attention on pollinator declines, there is still only a rudimentary understanding of the response of bees—the most important group of pollinators worldwide—to ongoing land use changes. Euglossine bees are an ecologically important Neotropical clade of forest-dependent pollinators. Despite the fact that euglossines are well studied relative to other groups of tropical bees, only three previous studies, all from Brazil, address the response of euglossines to forest fragmentation. In this study, I tripled the maximum sample size of previous efforts by sampling male euglossines in 22 forest fragments ranging in area from 0.25 ha to 230 ha in southern Costa Rica, using chemically baited Van Someren traps. Abundance of euglossine bees was significantly positively related to forest fragment size, negatively related to shape (edge:area ratio), and marginally related to fragment isolation. Euglossine species richness showed similar, but weaker trends: richness was significantly positively related to the quantity of forest edge, marginally negatively related to fragment area, and not related to fragment isolation. The positive relationship between euglossine richness and abundance and forest fragment edge is consistent with other studies that have found high euglossine density in secondary or disturbed forest. The data suggest that individual euglossines move between forest fragments, as has been shown in other systems. Still, forest fragmentation appears to affect euglossine bees more strongly than other bee groups in the study region. Their large flight range and positive relationship with forest edges may help to buffer the negative effects of fragmentation, allowing euglossines to utilize even the very smallest forest fragments in the study area.     

Effects of fragmentation on pollinator assemblage, pollen limitation and seed production of Mediterranean myrtle (Myrtus communis)

Pollinator assemblages may shift as a consequence of the destruction and fragmentation of natural habitats. The scarcity of mates and pollinators can lead plant populations to suffer from pollen limitation and a decrease in reproductive performance within fragmented areas. We studied the shift in pollinator assemblages along with pollen limitation and seed production patterns in the Mediterranean shrub Myrtus communis. Our study included six populations contrasting in patch and population size (Large vs. Small) within a fragmented landscape characterized by ∼1% of potential forest coverage. The breeding system in Myrtus communis was self-compatible, but compared with natural pollination, fruit set increased with pollen addition (quantity limited), and seed set (brood size) increased with outcross pollen addition (quality limited). While the pollinator assemblage in Large patches was taxonomically diverse, it was almost monopolized by honeybees in Small patches, where visitation rates were highest and wild bee species were almost absent. In general, Small populations were less pollen limited for fruit set than Large populations, particularly those that received the highest rates of honeybee visits. However, despite differences in fragmentation and pollinators between Large and Small populations, seed production patterns (brood size and seed mass) were rather similar among them, in agreement with similar pollen limitation levels found for brood size. A higher susceptibility of native pollinators to the presence of honeybee hives was found in Small patches, suggesting that the pollinator assemblage may be severely altered when fragmentation occurs in combination with beekeeping. We discuss its implications and effects on plant reproduction in fragmented areas.     

Effect of cultivation on microbial carbon and nitrogen in dry tropical forest soil

Summary Fifteen- and forty-year-old cropfields developed from a dry tropical forest were examined for soil organic C and total N and soil microbial C and N. The 15-year-old field had never been manured while the 40-year-old field had been fertilized with farmyard manure every year. The native forest soil was also examined. The results indicated that the native forest soil lost about 57% and 62% organic C and total N, respectively, in the 0–10 cm layer after 15 years of cultivation. The microbial C and N contents of the forest soil were greater than those of the cultivated soils. Application of farmyard manure increased the biomass-C and -N levels in the cultivated soil but the values were still markedly lower than in the forest soil. There was an appreciable seasonal variation in biomass C and N, the values being highest in summer and lowest in the rainy season. During an annual cycle, biomass-C contents varied from 180 to 727 g g^–1 and N from 20 to 80 g g^–1 dry soil, and both were linearly related. Microbial biomass C represented 1.6%–3.6% of total soil organic C and microbial biomass N represented 1.7% 1–4.4% of soil organic N.     

Depression of tree recruitment by the Pacific rat (Rattus exulans Peale) on New Zealand's northern offshore islands

Recruitment of tree populations on islands with and without Rattus exulans Peale, and recruitment changes on islands before and after rat eradication are compared using size-class analyses. Seedling establishment in rat-proof exclosures was compared with adjacent control areas. R. exulans depress recruitment of 11 species of coastal trees among 17 species studied: Coprosma macrocarpa, Coprosma repens, Dysoxylum spectabile, Melicytus novae-zelandiae, Nestegis apetala, Pisonia brunoniana, Pittosporum crassifolium, Pouteria costata, Pseudopanax lessonii, Rhopalostylis sapida and Streblus banksii; some to the point of local extinction. Where R. exulans has reached small islands supporting dense colonies of petrels (Procellariidae, Pelecanoididae), the numbers of some plant and animal species have been greatly reduced. Successional pathways in vegetation recovering on larger islands formerly occupied by Maori, have also been altered by this rat. Similar effects by R. exulans on the plant life of Pacific islands are probable considering the length of time R. exulans has been present.     

Forest fragment size and microhabitat effects on palm seed predation

The establishment of plant species depends crucially on where the seeds are deposited. However, since most studies have been conducted in continuous forests, not much is known about the effects of forest fragmentation on the maintenance of abiotic and biotic characteristics in microhabitats and their effects on seed survival. In this study, we evaluated the effects of forest fragmentation on the predation upon the seeds of the palm Syagrus romanzoffiana in three microhabitats (interior forest, forest edge and gaps) in eight fragments of semi-deciduous Atlantic forest ranging in size from 9.5 ha to 33,845 ha in southeastern Brazil. Specifically, we examined the influence of the microhabitat structure, fauna and fragment size on the pattern of seed predation. Fragments <100 ha showed similar abiotic and biotic characteristics to those of the forest edge, with no seed predation in these areas. Forest fragments 230-380 ha in size did not present “safe sites” for S. romanzoffiana seed survival and showed high seed predation intensity in all microhabitats evaluated. In fragments larger than 1000 ha, the seed predation was lower, with abiotic and biotic differences among gaps, interior forests and forest edges. In these fragments, the survival of S. romanzoffiana seeds was related to squirrel abundance and interior forest maintenance. Based on these results, we concluded that there are no safe sites for S. romanzoffiana seed establishment in medium- and small-sized fragments as result of the biotic and abiotic pressure, respectively. We suggest that on these forest fragments, management plans are needed for the establishment of S. romanzoffiana, such as interior forest improvement and development in small-sized sites in order to minimize the edge effects, and on medium-sized fragments, we suggest post-dispersal seed protection in order to avoid seed predation by vertebrates. Our findings also stress the importance of assessing the influence of forest fragmentation on angiosperm reproductive biology as part of the effective planning for the management of fragmented areas.     

Assessing landscapes: a case study of tree and shrub diversity in the seasonally dry tropical forests of Oaxaca, Mexico and southern Honduras

Tropical dry forests, with their distinct and economically important diversity, are acknowledged conservation priorities because of alarming rates of forest conversion. Whilst it is realised that terrestrial conservation requires an understanding of landscape level patterns of diversity, forests are rarely assessed accordingly. Here we demonstrate that, in the case of the seasonally dry tropical forests of the Pacific watershed of Mesoamerica, landscape level assessment of woody diversity can inform decision making relevant to both between-landscape and within-landscape prioritisation. We report floristic surveys of dry forest landscapes in Oaxaca, Mexico and southern Honduras. It is noted that these forests are floristically similar to other seasonally dry tropical forests in the neotropics. By calculation of Genetic Heat Indices, a relative measure of the concentration of restricted range species in a sample, we determine that the conservation of the tree diversity of the coastal lowlands of Oaxaca should be prioritised over that of southern Honduras. The current conservation status of forested areas in Oaxaca is briefly described. We suggest that the greater degree of anthropogenic disturbance in southern Honduras may explain the relative lack of restricted range species there. We argue that some forest fallows can act as analogues of mature forest and therefore landscape elements other than mature forest need to be included in forest conservation assessments. We conclude that diversity sampling of any forest type should not be limited to mature forests, but extended to other elements of forested landscapes.     

Effects of forest fragmentation on the granivory of differently sized seeds

The Maulino forest is a unique temperate ecosystem restricted to a small range of the coast of central Chile. This forest harbors many endemic species, and is threatened due to intensive deforestation and fragmentation. Currently the Maulino forest is composed of a suite of small fragments scattered in a landscape dominated by exotic plantations. The fragmentation of the Maulino forest has resulted in a higher abundance of granivores in small forest fragments compared with continuous forest. In order to determine if fragmentation-induced changes in granivore abundance affects the granivory of different size seeds, we experimentally assessed seed predation of a large-seeded species [Nothofagus glauca (Phil.) Krasser] and a small-seeded species [Nothofagus obliqua (Mirbel) Oersted] in the edges and interior of one continuous (large) forest and three small fragments (∼3 ha) surrounded by plantations of the exotic tree Pinus radiata. To determine what kind of granivores are preying upon seeds, seeds of both species were excluded from and exposed to large and small granivores. Granivory was higher in small fragments than in continuous forest, higher in the edges than in the forest interior, and higher upon large than on small seeds. Rodents, which were more abundant in forest fragments, were the main consumers. Thus, fragmentation indeed affects granivory increasing the consumption of seeds by predators inhabiting the Maulino forest remnants or coming from the matrix. This change may affect the future structure of the tree community in forest fragments.     

Seasonal changes in multi-scale spatial structure of soil pH and related parameters along a tropical dry evergreen forest slope

Seasonal changes in multi-scale spatial variation in soil chemical properties, which may be controlled simultaneously by biotic and abiotic factors, have not been studied in tropical dry forests. We evaluated the spatial variation of physico-chemical soil properties, plant litter and terrain attributes at multiple scales in a tropical dry evergreen forest using multivariate geostatistics. Soil samples were collected at different depths using nested interval sampling during 1- and 10-m intervals in both the wet and dry seasons. We measured pH, exchangeable cations (Ex-K⁺ and Ex-Ca²⁺), acidity (Ex-H⁺ and Ex-Al³⁺), particle size (clay and sand contents), and forest floor mass (O_i and O_a). Pronounced spatial variation in pH was observed in surface soil (0-5 cm) but not in deeper soil (5-55 cm). Multi-scale spatial structures with short (20 m) and long (86 m) ranges were observed in the auto- and cross-variograms of soil, litter and slope gradient. Pronounced multi-scale structures were observed simultaneously in pH and Ex-Ca²⁺ both in the wet and dry seasons. Only a short-range structure was observed in Ex-K⁺ and O_a, whereas a long-range structure was pronounced in sand contents and slope gradients. Although the variograms had similar shapes between wet and dry seasons for almost all variables, the short-range structure of the cross-variogram between O_a with pH and base cations was more pronouncedly developed in the wet season than in the dry season. Scale-dependent correlation coefficients suggest that a small-scale spatial variation in pH was connected to heterogeneous litter accumulation via base-cation input, whereas long-range spatial variation was simultaneously linked to particle size and slope gradient. This multivariate geostatistical approach applied within a stand detected biotic and abiotic factors controlling spatial variation in soil properties at both short and long distances.     

Abiotic and vertebrate seed dispersal in the Brazilian Atlantic forest: implications for forest regeneration

This study documents the proportion of woody plant species having different modes of seed dispersal, and classes of seed and fruit size at 16 Atlantic montane forest plots of southeastern Brazil (23-24° S). These plots represent six chronosequences (from 5-year-old to old-growth forest) of forest regeneration following small-scale shifting agriculture. Our results indicate that there is a gradual but predictable increase in the number of woody plant species relying on vertebrate-mediated seed dispersal in increasingly older plots. Moreover, the percentage of plant species bearing small seeds and fruits (< 0.6 cm in length) was more than halved from earlier to older forest plots, while the percentage of plant species producing middle-sized seeds and fruits at least doubled. Plant species belonging to the Melastomataceae, Myrsinaceae, Rubiaceae and Flacourtiaceae comprised a large proportion of the small-seeded and fruit species (33-53%), and largely occurred in early successional forest plots (5-30 years old). In contrast, species belonging to the Myrtaceae and Lauraceae represented most of the species (51-72%) bearing medium-sized seeds and fruits (0.6-1.5 cm in length), and occurred primarily in the old-growth forests. Shifts in the relative importance of dispersal strategies during the regeneration process of Atlantic montane forest, as well as in diaspore size appear to be related to the balance between early successional and shade-tolerant species associated with particular life forms and plant families. Finally, we discuss the relationships between the species richness of Myrtaceae and Lauraceae trees, seed dispersal by medium to large vertebrates, and possible regeneration scenarios for the Atlantic forest.     

A variable edge effect on trees of Bwindi Impenetrable National Park, Uganda, and its bearing on measurement parameters

With tropical forests under increased pressure from fragmentation and nascent effects of climate change, it is imperative to reliably predict change for timely reduction of impacts. This depends on measuring the right variables and analyzing them in ways that best detect change. I compared the usefulness of density, species richness, species diversity, and size in detecting edge penetration; and other parameters in revealing edge effect on tree species in Bwindi Impenetrable National Park, Uganda. Trees were sampled in 104 1 km edge-interior transects set around the entire park. Density and species richness were better indicators of edge-width than girth size and diversity measurements, both showing edge penetration as most intense within 300 m, although effect was detectable up to 900-1000 m. This estimate compared closely with an earlier estimate of 300-350 m obtained using incidences of resource harvest. Among other measures, measures of tree species abundance and frequency showed significant response to the edge by both early succession and primary forest species; while analyses of density showed edge response primarily by forest interior species. Parameters of tree size and their derivatives showed edge response by the lowest number of species. These results suggest that to measure edge effects reliably, it is important to select the right parameters.     

Mixed-species bird flocks from Brazilian Atlantic forest: the effects of forest fragmentation and seasonality on their size, richness and stability

The effects of forest fragmentation on species richness, size and stability of bird mixed-species flocks were studied during rainy and dry seasons at nine Atlantic Forest fragments in Minas Gerais state, southeastern Brazil. Two distinct kinds of mixed-species flocks were identified at the study area: heterogeneous flock and understory flocks. The first type of association was observed in all forest fragments and was composed mainly of canopy and midstory species, but may include a few understory species. Understory flocks were composed mainly of understory species and were detected in only three forest fragments. Forest fragment area and season significantly affected species richness, size and stability of heterogeneous flocks. Moreover, species richness in heterogeneous flocks was correlated with total species pool in forest fragments. Species richness and size of understory flocks were different among the three forest fragments, but season influenced only flock size. Understory flock stability differed neither among forest fragments nor between seasons. The red-crowned ant-tanager Habia rubica was the nuclear and leader species of understory flocks, and determined the occurrence of this association in forest fragments. Our results show that forest fragmentation can affect species’ interactions in foraging associations, and that conservation plans for the region should preserve nuclear species of mixed-species flocks and their preferred habitats.     

Habitat structure and proximity to forest edge affect the abundance and distribution of forest-dependent birds in tropical coastal forests of southeastern Madagascar

Despite the fact that Madagascar is classified a biological `hotspot' due to having both high levels of species endemism and high forest loss, there has been no published research on how Madagascan bird species respond to the creation of a forest edge or to degradation of their habitat. In this study, we examined how forest bird communities and different foraging guilds were affected by patch habitat quality and landscape context (forest core, forest edge and matrix habitat) in the threatened littoral forests of coastal southeastern Madagascar. We quantified habitat use and community composition of birds by conducting 20 point counts in each landscape contextual element in October and November 2002. We found that littoral forest core habitats had significantly (p<0.01) more bird species than forest edge and matrix habitats. Thirty-one (68%) forest dependent species were found to be edge-sensitive. Forest edge sites had fewer species, and a higher representation of common species than forest interior sites. Twenty-nine species were found in the matrix habitat, and the majority of matrix-tolerant forest species had their greatest abundance within littoral forest edge habitats. Guild composition also changed with landscape context. Unlike other tropical studies with which we are familiar, we found that frugivorous species were edge-sensitive while sallying insectivores were edge-preferring. The majority of canopy insectivores (n=15, 88%), including all six endemic vanga species, were edge-sensitive. When habitat quality was assessed, the distributions of nine edge-sensitive species were significantly (p<0.01) affected by changes in habitat complexity and vegetation vertical structure in core or edge point counts. Therefore, we believe that changes in vegetation structure at the edge of littoral forest remnants may be a key indicator of mechanisms involved in edge sensitivity of forest dependent species in these forests. Our findings indicate that habitat fragmentation and degradation affect Madagascan bird communities and that these processes threaten many species. With continued deforestation and habitat degradation in Madagascar, we predict the further decline of many bird species.     

Edge effects on recruitment of Trillium camschatcense in small forest fragments

To evaluate edge effects on recruitment of the understory perennial herb Trillium camschatcense, spatial distribution and stage-class structure based on four life-history stages were investigated within a small forest fragment in eastern Hokkaido, Japan. Seedlings were most affected by edge effects. The density of seedlings was considerably lower in the forest edges than in the interiors and was higher on the northern side of the forest. Although three stages suffered edge and orientation effects, flowering plants experienced much more moderate edge effects than juvenile stages and showed relatively uniform distribution. The results suggest that the recruitment of juvenile stages was strongly limited near forest edges. Seedling density was correlated with microclimatic conditions, not with seed production. The edge and orientation effects on stage-class structures were also observed in two other small populations, emphasizing the role of edge-related decreases in seed germination and subsequent survival for limiting the recruitment of small, fragmented populations of T. camschatcense.     

Bird species abundance–occupancy patterns and sensitivity to forest fragmentation: Implications for conservation in the Brazilian Atlantic forest

Developing a predictive theory for species responses to habitat fragmentation is a large, complex challenge in conservation biology, and meeting this challenge likely requires tailoring predictions to specific habitats and taxa. We evaluate the effects of fragmentation on forest birds living in three distinct forest ecosystems found in Brazilian Atlantic forest: seasonal semi-deciduous forest (SF), mixed rain forest (MF), and dense rain forest (DF). We test the hypotheses that (1) bird species most prevalent in SF (relative to other habitat types) will be least vulnerable to population declines in fragmented SF, and (2) species with stronger affiliations with DF or MF will be relatively more sensitive to fragmentation in SF. Using an exploratory statistical technique called “Rank Occupancy–Abundance Profiles (ROAPs),” we compared distribution and abundance of birds among large “continuous” areas of each forest type, then compared abundances in continuous SF forests with patterns of abundance in small fragments of SF, where edge effects could play a marked role in population dynamics. Overall, 39 species showed substantially lower local abundance, occupancy, or both in SF fragments versus continuous SF. As predicted, a higher proportion of bird species associated with DF appeared sensitive to fragmentation in SF; by contrast, species most abundant in SF and MF were similarly abundant in fragmented SF. Our study demonstrates how quantifying distribution and abundance in diverse habitats may enhance managers’ ability to incorporate species-specific responses to human disturbances in their conservation plans, and points out ways that even small reserves may have significant conservation value.     

The responses of understorey birds to forest fragmentation, logging and wildfires: An Amazonian synthesis

We combine mist-net data from 24 disturbance treatments taken from seven studies on the responses of understorey Amazonian birds to selective logging, single and recurrent wildfires, and habitat fragmentation. The different disturbance treatments had distinct effects on avian guild structure, and fire disturbance and the isolation of forest patches resulted in bird communities that were most divergent from those in continuous, undisturbed forest in terms of their species composition. Although low-intensity logging treatments had the least noticeable effects, the composition of understorey birds was still markedly different from the composition in undisturbed forest. This analysis demonstrates the importance of preventing habitat fragmentation and the spread of fires in humid tropical forests, and highlights the need for more research to determine the long-term suitability of large areas of degraded forest for forest birds.     

Dynamics of soil biomass C,N, and P in a dry tropical forest in India

Summary Three dry tropical forest soils along a topographic sequence were examined to determine the seasonal dynamics of microbial C, N, and P. The lowest microbial biomass was found in forest soils at the foot of the hill followed by midslope forest soils. The hilltop soil, which had the most fine particles, water-holding capacity, organic C, and total N, reflected the presence of greater amounts of microbial C, N, and P. Mean annual microbial C, N, and P ranges were 466–662, 48–72 to 21–30 g g^-1, respectively. The seasonal pattern of microbial biomass, C, N, and P was similar at all sites, the values being greatest during the dry season and lowest during the wet season. The seasonal values for microbial biomass C, N, and P were positively correlated with each other and a negative correlation was found between microbial biomass and the fine root mass in these forest soils.