Controlled experiments of habitat fragmentation: a simple computer simulation and a test using small mammals

Habitat fragmentation involves a reduction in the effective area available to a population and the imposition of hard patch edges. Studies seeking to measure effects of habitat fragmentation have compared populations in fragments of different size to estimate and area effect but few have examined the effect of converting open populations to closed ones (an effect of edges). To do so requires a shift in spatial scope-from comparison of individual fragments to that of fragmented versus unfragmented landscapes. Here we note that large-scale, controlled studies of habitat fragmentation have rarely been performed and are needed. In making our case we develop a simple computer simulation model based on how individual animals with home ranges are affected by the imposition of habitat edges, and use it to predict population-level responses to habitat fragmentation. We then compare predictions of the model with results from a field experiment on Peromyscus and Microtus. Our model treats the case where home ranges/territories fall entirely within or partially overlap with that of sample areas in continuous landscapes, but are restricted to areas within habitat fragments in impacted landscapes. Results of the simulations demonstrate that the imposition of hard edges can produce different population abundances for similar-sized areas in continuous and fragmented landscapes. This edge effect is disproportionately greater in small than large fragments and for species with larger than smaller home ranges. These predictions were generally supported by our field experiment. We argue that large-scale studies of habitat fragmentation are sorely needed, and that control-experiment contrasts of fragmented and unfragmented microlandscapes provide a logical starting point.     

Effect of fragmentation on the community structure of epixylic bryophytes in Atlantic Forest remnants in the Northeast of Brazil

The effects of fragmentation and edge effects on the floristic composition, richness, diversity and abundance of epixylic bryophytes (growing on decaying wood) were investigated in ten fragments of Atlantic Forest remnants in the Northeast of Brazil. In each fragment, four perpendicular 100 m transects were demarcated. Along these transects, samples of bryophytes growing on decaying wood were collected. The forest fragments were grouped in three size classes (small: <100 ha; medium: 100–500 ha; large: >500 ha). Correlation and multivariate analysis were undertaken between bryophyte flora and fragment metrics (size, form, isolation, altitude variation, nuclear area and secondary vegetation percentage and distance from the edge). A total of 99 species of bryophytes, 52 liverworts and 47 mosses were registered. The statistical results confirming fragment size is an important factor in epixylic community structure. Therefore, composition, richness, diversity and abundance can be better explained by a junction of all studied landscape factors. Bryophyte richness, the percentage of samples with the greatest coverage of decaying wood and shade-tolerant species’ distribution, were not correlated to distance from the forest edge. This suggests that edge effects are not linear or can be detected beyond 100 m from the edge, which is very important for inclusion in future studies.     

The historical bridge between the Amazon and the Atlantic Forest of Brazil: a study of molecular phylogeography with small mammals

Abstract Aim To examine how the genetic diversity of selected taxa of forest‐dwelling small mammals is distributed between and within the major rain forest domains of Amazonia and Atlantic Forest and the intervening interior forests of Brazil, as inferred by the relationships between gene genealogies and geography. I also addressed the historical importance of the central Brazilian forests in connecting Amazon and Atlantic Forest populations of rodents and marsupials. Methods I evaluated variation in the mitochondrial cytochrome b gene to estimate the levels of sequence divergence between those taxa occurring throughout the Amazon, Atlantic Forest, and forests in the Cerrado and Caatinga regions. I inferred the hierarchical relationships between haplotypes, populations and formal taxa using the cladistic approach of maximum parsimony. I compared areas and the clades identified by superimposing cladograms on the geographical distribution of samples. The degree of concordance both in phylogeny and the depth of the nodes in these phylogenies, in addition to patterns of geographical distribution of clades, permitted me to make inferences on how, when and where the taxa differentiated. Results Sequence similarity is often greater between samples from the Atlantic Forest and either Amazon or central Brazilian forests than it is within each of the two rain forest domains. The Atlantic Forest clades are either not reciprocally monophyletic or are the sister group to all the other clades. There is some indication of northern and southern components in the Atlantic Forest. Given the geographical distribution of clades and the relatively deep levels of divergence, the central Brazilian area does not behave as a separate region but is complementary to either Amazon or Atlantic Forest. Patterns of area relationships differ across taxa, suggesting that different processes and/or historic events affected the diversification within each lineage. Main conclusions The Amazon and the Atlantic forests are not exclusive in terms of their small mammal faunas; both overlap broadly with taxa occurring in gallery forests and dry forests in central Brazil. Central Brazilian forests are an integral part of the evolutionary scenario of lowland small mammals, playing an important role as present and past habitats for rain forest species. Therefore, representatives from this area should always be included in analyses of the evolutionary history of lowland rain forest faunas. The incongruence of branching patterns among areas is in agreement with recent results presented for Neotropical passerine birds and indicates that a single hypothesis of Neotropical area relationships is unlikely. These findings reinforce the idea that speciation in the Neotropics will not be explained by any single model of vicariance or climatic changes.     

Effects of fragmentation on forest structure and litter dynamics in Atlantic rainforest in Pernambuco, Brazil

This study evaluates the effects of fragmentation on the spatial and temporal dynamics of small litterfall production in Atlantic rainforest in Pernambuco State, Brazil. Litterfall was collected for 24 months at two 0.2 ha sites, located in the forest edge zone and the forest interior, within a rainforest patch of about 300 ha. Structural parameters of both forest sites were recorded. Litter was sorted into six fractions (foliage, twigs, buds/flowers, fruits/seeds, peduncles, rest), dried and weighed. The interior forest plot contained 314 live trees with a dbh 5 cm and a stand basal area of 41.8±8.7 m², whereas the forest edge contained 211 live trees and a stand basal area of 23.4±3.6 m². Total small litterfall was extraordinarily high and totalled 12.62±4.73 t ha⁻¹ yr⁻¹ in interior forest and 14.74±2.78 t ha⁻¹ yr⁻¹ in forest edge. High litterfall rates are probably due to a pronounced periodicity, edge effects alter litterfall strongly.     

Linking changes in small mammal communities to ecosystem functions in an agricultural landscape

Global increases in agricultural production have significant implications for biodiversity and ecosystem processes. In southern Africa, sugarcane production has converted native vegetation into agricultural monocultures. We examined functional group abundance along a conservation-agriculture gradient in the Lowveld of Swaziland. We captured small mammals representing 4 functional groups: omnivores, insectivores, granivores, and herbivores and found evidence of distinct changes in small mammal functional groups across the conservation-agriculture boundary. Granivores declined with increasing distance into the sugarcane and were completely absent at 375 m from the boundary while omnivores increased in the sugarcane. Insectivores and herbivores showed no differences between the two land uses; however, during the dry season, there were significantly more insectivores at the conservation-agriculture interface than in the conservation lands. Shifts in small mammal communities have clear implications for ecosystem processes as the removal of granivores from savannah systems can drastically alter vegetative structure and potentially lead to shrub encroachment via reduced levels of seed predation, while abundant omnivorous small mammals can cause significant crop damage and increase the prevalence of vector borne diseases in the environment.     

Differences in seed rain composition in small and large fragments in the northeast Brazilian Atlantic Forest

Tropical forests are seriously threatened by fragmentation and habitat loss. The impact of fragment size and forest configuration on the composition of seed rain is insufficiently studied. For the present study, seed rain composition of small and large forest fragments (8–388 ha) was assessed in order to identify variations in seed abundance, species richness, seed size and dispersal mode. Seed rain was documented during a 1‐year period in three large and four small Atlantic Forest fragments that are isolated by a sugarcane matrix. Total seed rain included 20,518 seeds of 149 species of trees, shrubs, palms, lianas and herbs. Most species and seeds were animal‐dispersed. A significant difference in the proportion of seeds and species within different categories of seed size was found between small and large fragments. Small fragments received significantly more very small‐sized seeds (<0.3 cm) and less large‐seeded species (>1.5 cm) that were generally very rare, with only one species in small and eight in large fragments. We found a negative correlation between the inflow of small‐sized seeds and the percentage of forest cover. Species richness was lower in small than in large fragments, but the difference was not very pronounced. Given our results, we propose changing plant species pools through logging, tree mortality and a high inflow of pioneer species and lianas, especially in small forest fragments and areas with low forest cover. Connecting forest fragments through corridors and reforestation with local large‐seeded tree species may facilitate the maintenance of species diversity.     

Effects of fragmentation on parasite burden (nematodes) of generalist and specialist small mammal species in secondary forest fragments of the coastal Atlantic Forest,Brazil

Parasites are considered to play an important role in the regulation of wild animal populations. We investigated parasite burden of gastrointestinal nematodes and body condition in specialist and generalist small mammal species in secondary forest fragments in the highly endangered coastal Atlantic Forest. We hypothesized that body condition decreases with increasing parasite load and that parasite burden increases with increasing fragmentation in specialist species but not in generalist species as a consequence of differing responses to fragmentation effects. Investigated species were Akodon montensis, Oligoryzomys nigripes, and Delomys sublineatus (rodents) and the marsupials Marmosops incanus and Gracilinanus microtarsus. Prevalence of parasites was high in all species except for the arboreal G. microtarsus, presumably because of decreased infection probability. No correlation was found between body condition and parasite load in any of the species. Contrary to our expectations, body condition of the specialists D. sublineatus and M. incanus increased in both species with increasing fragmentation. In D. sublineatus, parasite burden increased and body condition decreased in fragments with relatively high density probably due to increased contact rates and facilitation of infection with nematodes. In all generalist species, low or no correlation between parasite burden and fragmentation was detected, suggesting little effect of fragmentation on population health.     

长白山森林景观破碎的遥感探测

天然林数量的减少以及质量的降低对中国的生态环境和经济发展产生不利影响。虽然林地的破碎对各种自然灾害起到关键性作用,林地是如何被破碎的尚不十分清楚。应用卫星遥感技术,以及地理信息系统和空间分析手段研究中国东北－原始的针阔混交林的破碎过程。结果表明,本来完整的原始森林,已经被一种“复叶”状小面积皆伐作业分割为零碎的斑真。采伐地的面积平均为15hm^2, 但随着采伐的继续、采伐地段连为一体,增大了采伐迹地的面积,从而保留的林地面积越来越小。采伐地点的选择没有考虑物种和环境的保护,林地破碎的特征是保留林地面积的缩小,中、小林地数量的增加,林地斑块形状的改变,以及林缘长度的增加。最后就保护东北森林植被完整性的角度提出林业政策与森林经营的改进意见。     

Differences in mammal communities between forest fragments and restoration areas in the Atlantic Forest

The persistence of larger mammals in fragmented forest landscapes depends not only on the protection of remaining habitats but also on ecological restoration sites. It is known that the landscape context is an important predictor of species persistence, abundance and distribution. Here we evaluate how landscape characteristics influence the recovery of larger mammals in ecological restoration sites. We assess the richness and composition of mammals in forest fragments and restoration sites using landscape metrics such as forest cover and connectivity. Forest fragments and restoration sites present the same richness (n = 26), but differ in species composition. Some seed-dispersing mammals were absent in restoration areas, such as Alouatta guariba (brown howler monkey) and Coendou spinosus (paraguayan hairy dwarf porcupine). The percentage of forest cover in the landscape was responsible for 29.09% of the variation in species composition between the evaluated forest formations, exerting a positive or negative influence depending on the species requirements. The results demonstrate the importance of considering not only landscape metrics in an ecological restoration plan, but also the historical landscape context, such as the fauna composition before the disturbance and how these species respond to environmental changes, thus improving the success of future ecological restoration measurements and policies.     

Ecological fragmentation effects in mouse lemurs and small mammals in northwestern Madagascar

Habitat loss and fragmentation are major ecological forces threatening animal communities across the globe. These issues are especially true in Madagascar, where forest loss is ongoing. We examined the effects of forest fragmentation on the distribution and abundance of sympatric, endemic gray, and golden-brown mouse lemurs (Microcebus murinus and Microcebus ravelobensis), the endemic western tuft-tailed rat (Eliurus myoxinus), and the invasive black rat (Rattus rattus) in two regions in northwestern Madagascar. We used systematic capture procedures in 40 forest fragments and four continuous forest sites which differed in size, shape, and degree of isolation. With a trapping effort of 11,567 trap nights during two dry seasons (2017–2018), we captured 929 individuals (432 M. ravelobensis, 196 M. murinus, 116 E. myoxinus, and 185 R. rattus). We examined the influence of study region, forest type (fragment vs. continuous), forest size, forest shape, the proportion of 50-m forest edge and distance to continuous forest on the abundance and interaction of the four species. Responses to fragmentation differed strongly between species, but no interaction could be detected between the abundance of the different species. Thus competition within and between native and invasive species may not be regulating abundances in these regions. On the contrary, the abundance of M. ravelobensis and E. myoxinus differed significantly between study regions and was negatively affected by fragmentation. In contrast, there was no evidence of an impact of fragmentation on the abundance of M. murinus. Finally, the invasive R. rattus responded positively to the increasing distance to the continuous forest. In conclusion, the response of small Malagasy mammals to forest fragmentation varies largely between species, and fragmentation effects need to be examined at a species-specific level to fully understand their ecological dynamics and complexity.     

Strong influence of long-distance edge effect on herb-layer vegetation in forest fragments in an agricultural landscape

In agricultural landscapes in central Europe, species richness of the herbaceous plant community may be compromised by processes associated with forest fragmentation, habitat loss, and management practices. We examined variability in species richness and composition of the herbaceous layer in 229 plots located in 23 forest fragments (0.1 to 255 ha), in a representative upland agricultural landscape in central Bohemia, in relation to the most important site environmental factors, edge effects, and site history. The influence of environmental factors on the composition of vegetation in the herb layer was evaluated using generalized additive models, which enabled us to analyze highly non-linear and non-monotonic relationships. Total species richness and number of red-listed and ancient forest species were significantly influenced by type of forest vegetation, light quality, soil pH, slope aspect, and distance from the forest edge. Implications of the significant explanatory variables corresponded well to previous findings, with the exception of distance from the forest edge, for which we found a positive relationship with species richness for distances up to 200 m toward the forest interior. Plant species with low colonization ability occupied plots with increasing frequency from edge to forest interior, while fast-colonizing species showed the opposite trend. Apart from the edge effect, forest continuity should be considered for its important contribution to the richness of ancient forest and red-listed species, whereas the effect of forest fragment size appeared to be generally weak. These results do not negate the importance of large forest fragments for the maintenance of herb layer species richness, but specifically emphasize the essential contribution of the core habitats of these forests. In summary, we showed that the negative effects of habitat fragmentation on the richness of ancient forest and red-listed species and on herb layer species in total can be largely attributed to either the edge effect itself or to aggregate effects of forest edge and forest continuity.     

Forest fragmentation drives Atlantic forest of northeastern Brazil to biotic homogenization

Aim To examine whether the tree flora of the Atlantic forest of northeastern Brazil has experienced detectable taxonomic homogenization via the proliferation of native pioneer species in response to habitat loss and fragmentation. Location Biotic homogenization (BH) was examined across the Atlantic forest of northeast Brazil, i.e. a 56,000 km² piece of tropical forest and a distinct centre of species endemism in South America. Methods We assessed a dataset consisting of 5122 tree records and compared the similarity of tree floras from 12 semi‐natural sub‐regions of the Atlantic forest between two time periods: pre‐1980 (plant records between 1902 and 1980), and post‐1980 (between 1981 and 2006). To understand the mechanisms leading to BH (1) tree floras were ordered (via non‐metric multidimensional scaling – NMDS) by date (pre/post 1980) based on species occurrence and frequency, (2) NMDS axes were regressed against the proportion of those species that increased their occurrence post‐1980 (i.e. the winner species), and (3) patterns of geographic distribution and frequency of particular life‐history traits were examined across winner species and a control group. Results Tree floras across the Atlantic forest became c. 20–40% more similar to each other post‐1980, but patterns of species similarity were also influenced by between‐plot geographical distance. NMDS ordination clearly segregated pre‐ and post‐1980 floras with a clear signal of floristic convergence. Furthermore, winner tree species were largely composed of short‐lived and small‐seeded pioneer species that exhibit wide geographic distributions. Main conclusions Our results suggest that tropical forest biotas are susceptible to taxonomic homogenization (i.e. increasing levels of similarity) in the context of severe human‐disturbance via the proliferation of particular groups of native species comprised mainly by ecologically‐plastic, generalist species. We are thus extending the concept of homogenization to address and highlight a pervasive biological shift in the structure of tropical forest communities currently taking place across hyper‐fragmented landscapes.     

Landscape attributes as drivers of the geographical variation in density of Sapajus nigritus Kerr, 1792, a primate endemic to the Atlantic Forest

Neotropical primates are among the most well studied forest mammals concerning their population densities. However, few studies have evaluated the factors that influence the spatial variation in the population density of primates, which limits the possibility of inferences towards this animal group, especially at the landscape-level. Here, we compiled density data of Sapajus nigritus from 21 forest patches of the Brazilian Atlantic Forest. We tested the effects of climatic variables (temperature, precipitation), landscape attributes (number of patches, mean inter-patch isolation distance, matrix modification index) and patch size on the population density using linear models and the Akaike information criterion. Our findings showed that the density of S. nigritus is influenced by landscape attributes, particularly by fragmentation and matrix modification. Overall, moderately fragmented landscapes and those surrounded by matrices with intermediate indexes of temporal modification (i.e., crop plantations, forestry) are related to high densities of this species. These results support the assumptions that ecologically flexible species respond positively to forest fragmentation. However, the non-linear relationship between S. nigritus density and number of patches suggests that even the species that are most tolerant to forest cover changes seem to respond positively only at an intermediate level of habitat fragmentation, being dependent of both a moderate degree of forest cover and a high quality matrix. The results we found here can be a common response to fragmentation for those forest dweller species that are able to use the matrix as complementary foraging sites.     

Diversity and habitat association of small mammals in Aridtsy forest,Awi Zone,Ethiopia

Here, we conducted a survey to examine the diversity, distribution and habitat association of small mammals from August 2011 to February 2012 incorporating both wet and dry seasons in Aridtsy forest, Awi Zone, Ethiopia. Using Sherman live traps and snap traps in four randomly selected trapping grids, namely, natural forest, bushland, grassland and farmland, a total of 468 individuals comprising eight species of small mammals(live traps) and 89 rodents of six species(snap traps) were trapped in 2352 and 1200 trap nights, respectively. The trapped small mammals included seven rodents and one insectivore: Lophuromys flavopuntatus(30.6%), Arvicanthis dembeensis(25.8%), Stenocephalemys albipes(20%), Mastomys natalensis(11.6%), Pelomys harringtoni(6.4%), Acomys cahirinus(4.3%), Lemniscomys zebra(0.2%) and the greater red musk shrew(Crocidura flavescens, 1.1%). Analysis showed statistically significant variations in the abundance and habitat preferences of small mammals between habitats during wet and dry seasons.     

Forest fragmentation relaxes natural nest predation in an Afromontane forest

Nest predation is widely regarded as a major driver underlying the population dynamics of small forest birds. Following forest fragmentation and the subsequent invasion by species from non-forested landscape matrices, shifts in predator communities may increase nest predation near forest edges. However, effects of human-driven habitat change on nest predation have mainly been inferred from studies with artificial nests, despite being regarded as poor surrogates for natural ones. We studied variation in predation rates, and relationships with timing of breeding and characteristics of microhabitats and fragments, on natural white-starred robin Pogonocichla stellata nests during three consecutive breeding seasons (2004–2007) in a Kenyan fragmented cloud forest. More than 70% of all initiated nests were predated during each breeding season. Predation rates nearly quadrupled between the earliest and the latest nests within a single breeding season, increased with distance to the forest edge, and decreased with the edge-to-area ratio of forest fragments. These spatial relationships oppose the traditional perception of edge and fragmentation effects on nest predation, but are in line with results from artificial nest experiments in other East African forests. In case of inverse edge and fragmentation effects on nest predation, such as shown in this study, species that tolerate edges for breeding may be affected positively, rather than negatively, by forest fragmentation, while the opposite can be expected for species restricted to the forest interior. The possibility of inverse edge effects, and its conservation implications, should therefore be taken into account when drafting habitat restoration plans.     

Response of wood-inhabiting fungal community to fragmentation in a beech forest landscape

Fragmentation of natural habitats has become one of the main causes of the loss of biodiversity. To assess the effects of forest fragmentation on wood-inhabiting fungal community in a beech-dominated landscape, 15 differently shaped beech forest fragments were examined in northern Spain. This work covers all the wood-inhabiting macromycetes, including Basidiomycota and Ascomycota. A modelling approach was used to examine the predictability of the fungal community in a fragmented beech forest landscape. In the beech forest patches, a large proportion of edge, low tree densities and low levels of variety of woody debris caused a decrease of wood-inhabiting fungal richness. The fungal community composition proved complex to model due to its specific traits: it is made up of many species, most of which are rare, and each fungal group responds differently to environmental variables. Nevertheless, the dead wood availability and the exposure to light significantly affected the fungal community composition.     

Bird Species Sensitivity in a Fragmented Landscape of the Atlantic Forest in Southern Brazil1

I tested whether terrestrial/understory habits, large body sizes, low tolerance to edge or to matrix habitat, and food/habitat specialization are characteristics of the most vulnerable bird species based on point counts conducted in 14 forest fragments in northern Paraná, southern region of the Brazilian Atlantic forest. In addition, the sensitivity level of each bird species to forest fragmentation in northern Paraná (high, medium, and low) was compared to the Viçosa region, another fragmented landscape of the Atlantic forest. Contrary to expectations, large body size and terrestrial/understory habits were not features clearly related to high sensitivity to forest fragmentation; but species with tolerance to edge did have lower sensitivity. I found also that specialized habitat/food were not good predictors of extinctions; the most specialized guilds, bamboo and vine‐tangle insectivores and trunk and twig insectivores, had a high number of species with low sensitivity. Patterns of sensitivity were compared for 75 bird species that occurred in both northern Paraná and Viçosa; higher incongruence occurred for 14 and 7 species considered as highly sensitive in Viçosa but were of medium or low sensitivity, respectively, in northern Paraná. These differences may occur because sensitivity to habitat fragmentation may vary from the center to the periphery of a species' geographic range. Geographic variation in patterns of sensitivity to forest fragmentation may be important for conservation planning.     

Ground nest predation might not be higher along edges of Neotropical forest remnants surrounded by pastures: evidence from the Brazilian Atlantic forest

Several studies indicate that nest predation is higher along edges than in habitat interiors mainly due generalist predators arising from or proliferating in the surrounding matrix. Recent reviews demonstrate however that this is far from universal, in part because studies are strongly biased in temperate regions. Far fewer are known from the Neotropics and just a handful of studies have been carried out in the biologically-rich but severely fragmented Atlantic Forest of Brazil. Here we tested the influence of edge proximity on ground nest predation in a large (21,787 ha) Atlantic forest reserve. The experiment was carried out using chicken and quail eggs in 12 transects with 500 m in length, half of which parallel to internal edges (dirt roads) and half parallel to external edges (forest/pasture). Nest predation was significantly higher in wet season (42.7%), when no difference was found between edge and forest interior, than in dry season (16.5%), when nest predation was higher in forest interior (400 m). Within seasons, the difference between internal and external edges and the association between edge distance and edge type were not significant. Results suggest that ground nest predation in large protected areas of the Atlantic forest is mainly caused by forest dwellers rather than by species inhabiting the surrounding grassland-dominated landscape, mirroring recent findings in other tropical areas.