Effects of the Surrounding Matrix on Tree Recruitment in Amazonian Forest Fragments

Abstract:  Little is known about how the surrounding modified matrix affects tree recruitment in fragmented forests. We contrasted effects of two different matrix types, Vismia - and Cecropia -dominated regrowth, on recruitment of pioneer tree species in forest fragments in central Amazonia. Our analyses were based on 22, 1-ha plots in seven experimental forest fragments ranging in size from 1 to 100 ha. By 13 to 17 years after fragmentation, the population density of pioneer trees was significantly higher in plots surrounded by Vismia regrowth than in plots surrounded by Cecropia regrowth, and the species composition and dominance of pioneers differed markedly between the two matrix types. Cecropia sciadophylla was the most abundant pioneer in fragments surrounded by Cecropia regrowth (constituting nearly 50% of all pioneer trees), whereas densities of species in Vismia -surrounded fragments were distributed more evenly. Thus the surrounding matrix had a strong influence on patterns of tree recruitment in Amazonian forest fragments.     

Effects of Pioneer Tree Species Hyperabundance on Forest Fragments in Northeastern Brazil

Abstract: Despite many studies on fragmentation of tropical forests, the extent to which plant and animal communities are altered in small, isolated forest fragments remains obscure if not controversial. We examined the hypothesis that fragmentation alters the relative abundance of tree species with different vegetative and reproductive traits. In a fragmented landscape (670 km²) of the Atlantic Forest of northeastern Brazil, we categorized 4056 trees of 182 species by leafing pattern, reproductive phenology, and morphology of seeds and fruit. We calculated relative abundance of traits in 50 1‐ha plots in three types of forest configurations: forest edges, small forest fragments (3.4–83.6 ha), and interior of the largest forest fragment (3500 ha, old growth). Although evergreen species were the most abundant across all configurations, forest edges and small fragments had more deciduous and semideciduous species than interior forest. Edges lacked supra‐annual flowering and fruiting species and had more species and stems with drupes and small seeds than small forest fragments and forest interior areas. In an ordination of species similarity and life‐history traits, the three types of configurations formed clearly segregated clusters. Furthermore, the differences in the taxonomic and functional (i.e., trait‐based) composition of tree assemblages we documented were driven primarily by the higher abundance of pioneer species in the forest edge and small forest fragments. Our work provides strong evidence that long‐term transitions in phenology and seed and fruit morphology of tree functional groups are occurring in fragmented tropical forests. Our results also suggest that edge‐induced shifts in tree assemblages of tropical forests can be larger than previously documented.     

Fragmentation drives tropical forest fragments to early successional states: A modelling study for Brazilian Atlantic forests

Land use leads to massive habitat destruction and fragmentation in tropical forests. Despite its global dimensions the effects of fragmentation on ecosystem dynamics are not well understood due to the complexity of the problem. We present a simulation analysis performed by the individual-based model FORMIND. The model was applied to the Brazilian Atlantic Forest, one of the world's biodiversity hot spots, at the Plateau of São Paulo. This study investigates the long-term effects of fragmentation processes on structure and dynamics of different sized remnant tropical forest fragments (1-100 ha) at community and plant functional type (PFT) level. We disentangle the interplay of single effects of different key fragmentation processes (edge mortality, increased mortality of large trees, local seed loss and external seed rain) using simulation experiments in a full factorial design.Our analysis reveals that particularly small forest fragments below 25 ha suffer substantial structural changes, biomass and biodiversity loss in the long term. At community level biomass is reduced up to 60%. Two thirds of the mid- and late-successional species groups, especially shade-tolerant (late successional climax) species groups are prone of extinction in small fragments. The shade-tolerant species groups were most strongly affected; its tree number was reduced more than 60% mainly by increased edge mortality. This process proved to be the most powerful of those investigated, explaining alone more than 80% of the changes observed for this group. External seed rain was able to compensate approximately 30% of the observed fragmentation effects for shade-tolerant species.Our results suggest that tropical forest fragments will suffer strong structural changes in the long term, leading to tree species impoverishment. They may reach a new equilibrium with a substantially reduced subset of the initial species pool, and are driven towards an earlier successional state. The natural regeneration potential of a landscape scattered with forest fragments appears to be limited, as external seed rain is not able to fully compensate for the observed fragmentation-induced changes. Our findings suggest basic recommendations for the management of fragmented tropical forest landscapes.     

Effect of Forest Fragmentation on Genetic Diversity and Mating System in a Tropical Tree, Pithecellobium elegans

Genetic diversity, population differentiation, and temporal variation in outcrossing rates were examined for Pithecellobium elegans , a Neotropical rain forest canopy tree. Several forest fragments and a large reserve (1500 ha) were compared for several population genetic parameters. For eight populations sampled on the Atlantic coastal plain of Costa Rica, allozyme heterozygosity (0.13), polymorphism (35%), and effective number of alleles (1.24) were similar to values reported for other tropical tree species that occur at similar densities of less than one individual per hectare. These measures of genetic variation were lowest in populations of the smallest size, farthest from the reserve, and more isolated from other populations. Differentiation among samples collected in small forest fragments and the reserve population accounted for 10% of the total genetic variation observed. There was a positive relationship between the level of differentiation of populations from the reserve population and their distance from the reserve. Though predominantly an annually flowering species, the number of trees in flower at any one time varied from 80% of observed trees to only 6%. Outcrossing rates did not differ for two episodes in which the proportions of flowering trees were 33% and 80%. But periods of low density of flowering adults resulted in poor seed crops or failure to set fruit for many individuals. Population size at many sites will be effectively decreased because of the variation in flowering. Fragmentation of what was once a large, continuous forested area is resulting in genetic erosion of small, isolated populations of Pithecellobium elegans .     

Rain forest fragmentation and the proliferation of successional trees

The effects of habitat fragmentation on diverse tropical tree communities are poorly understood. Over a 20-year period we monitored the density of 52 tree species in nine predominantly successional genera (Annona, Bellucia, Cecropia, Croton, Goupia, Jacaranda, Miconia, Pourouma, Vismia) in fragmented and continuous Amazonian forests. We also evaluated the relative importance of soil, topographic, forest dynamic, and landscape variables in explaining the abundance and species composition of successional trees. Data were collected within 66 permanent 1-ha plots within a large (approximately 1000 km2) experimental landscape, with forest fragments ranging from 1 to 100 ha in area. Prior to forest fragmentation, successional trees were uncommon, typically comprising 2-3% of all trees (> or =10 cm diameter at breast height [1.3 m above the ground surface]) in each plot. Following fragmentation, the density and basal area of successional trees increased rapidly. By 13-17 years after fragmentation, successional trees had tripled in abundance in fragment and edge plots and constituted more than a quarter of all trees in some plots. Fragment age had strong, positive effects on the density and basal area of successional trees, with no indication of a plateau in these variables, suggesting that successional species could become even more abundant in fragments over time. Nonetheless, the 52 species differed greatly in their responses to fragmentation and forest edges. Some disturbance-favoring pioneers (e.g., Cecropia sciadophylla, Vismia guianensis, V. amazonica, V. bemerguii, Miconia cf. crassinervia) increased by >1000% in density on edge plots, whereas over a third (19 of 52) of all species remained constant or declined in numbers. Species responses to fragmentation were effectively predicted by their median growth rate in nearby intact forest, suggesting that faster-growing species have a strong advantage in forest fragments. An ordination analysis revealed three main gradients in successional-species composition across our study area. Species gradients were most strongly influenced by the standlevel rate of tree mortality on each plot and by the number of nearby forest edges. Species-composition also varied significantly among different cattle ranches, which differed in their surrounding matrices and disturbance histories. These same variables were also the best predictors of total successional-tree abundance and species richness. Successional-tree assemblages in fragment interior plots (>150 m from edge), which are subjected to fragment area effects but not edge effects, did not differ significantly from those in intact forest, indicating that area effects per se had little influence on successional trees. Soils and topography also had little discernable effect on these species. Collectively, our results indicate that successional-tree species proliferate rapidly in fragmented Amazonian forests, largely as a result of chronically elevated tree mortality near forest edges and possibly an increased seed rain from successional plants growing in nearby degraded habitats. The proliferation of fast-growing successional trees and correlated decline of old-growth trees will have important effects on species composition, forest dynamics, carbon storage, and nutrient cycling in fragmented forests.     

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.     

Local Extinctions of Terrestrial Insectivorous Birds in a Fragmented Landscape near Manaus, Brazil

Abstract: We examined the distributions of nine species of terrestrial insectivorous birds in 4- to 14-year-old rainforest fragments north of Manaus, Brazil. We surveyed 11 fragments of 1, 10, and 100 ha, 95 ha of secondary vegetation, and nine continuous forest plots (controls) of 1–100 ha. We augmented standard spot-mapping with extensive playback surveys. The fragments had been sampled with mist nets before isolation, so our results could be compared with the pre-isolation distribution. For the nine species, there were 55 cases of local extinction in the 11 fragments between about 1 year after isolation and the time of our surveys. This corresponds to 74% extinction of the local populations in fragments. These extinctions occurred despite the second-growth connection of some fragments to continuous forest as little as 70 m away. Three apparent colonization events by species not detected before isolation also occurred, but these may also reflect inadequate sampling before isolation. Our comparison of fragments and similar-sized control plots in continuous forest showed an area effect on species richness in both fragments and control plots, but fragments had fewer species than control plots of equal size. In a fragmented Amazonian landscape, the full suite of terrestrial insectivores would persist in the short term only in large fragments ( > 100 ha), although much larger areas are probably necessary for the long-term persistence of their populations.     

The impact of fragmentation and density regulation on forest succession in the Atlantic rain forest

The Atlantic Rain Forest, an important biodiversity hot spot, has faced severe habitat loss since the last century which has resulted in a highly fragmented landscape with a large number of small forest patches (<100 ha). For conservation planning it is essential to understand how current and future forest regeneration depends on ecological processes, fragment size and the connection to the regional seed pool. We have investigated the following questions by applying the forest growth simulation model FORMIND to the situation of the Atlantic Forest in the state of São Paulo, SE Brazil: (1) which set of parameters describing the local regeneration and level of density regulation can reproduce the biomass distribution and stem density of an old growth forest in a reserve? (2) Which additional processes apart from those describing the dynamics of an old growth forest, drive forest succession of small isolated fragments? (3) Which role does external seed input play during succession? Therefore, more than 300 tree species have been classified into nine plant functional types (PFTs), which are characterized by maximum potential height and shade tolerance. We differentiate between two seed dispersal modes: (i) local dispersal, i.e. all seedlings originated from fertile trees within the simulated area and (ii) external seed rain. Local seed dispersal has been parameterized following the pattern oriented approach, using biomass estimates of old growth forest. We have found that moderate density regulation is essential to achieve coexistence for a broad range of regeneration parameters. Considering the expected uncertainty and variability in the regeneration processes it is important that the forest dynamics are robust to variations in the regeneration parameters. Furthermore, edge effects such as increased mortality at the border and external seed rain have been necessary to reproduce the patterns for small isolated fragments. Overall, simulated biomass is much lower in the fragments compared to the continuous forest, whereas shade tolerant species are affected most strongly by fragmentation. Our simulations can supplement empirical studies by extrapolating local knowledge on edge effects of fragments to larger temporal and spatial scales. In particular our results show the importance of external seed rain and therefore highlight the importance of structural connectivity between regenerating fragments and mature forest stands.     

Lemurs and the Regeneration of Dry Deciduous Forest in Madagascar

Abstract: We sought to assess the role of lemurs for seed dispersal in the dry deciduous forest of western Madagascar and the possible consequences of the demise of lemurs for forest regeneration. Forest regeneration was studied in eight plots in two large blocks of primary forest and in seven fragments of primary forest (1 plot per fragment). In 4 of the 15 study plots, the abundance of saplings was negatively and significantly correlated ( p < 0.05) with the abundance of mature individuals of the same tree species. In another 10 study plots there were negative correlations, although these were not significant on the community level. Second-order statistics were significant with p < 0.001 and indicated that seed dispersal away from the parent trees was important for successful establishment of saplings. Apart from possibly the bush pig ( Potamochoerus larvatus ), only one vertebrate species of the dry forest, the brown lemur ( Eulemur fulvus ), ingested seeds> 11 mm long and passed them through its digestive tract unharmed. These results for lemurs were based on direct observations and fecal analyses. To evaluate the role of E. fulvus , we compared regeneration in forest plots with and without E. fulvus . In forest fragments without E. fulvus , fewer lemur-dispersed tree species regenerated than would be expected based on the presence of mature tree species that are lemur-dispersed ( p < 0.05). No such effect was seen in primary forests with E. fulvus or for trees whose seeds can also be dispersed by other vertebrates. Thus, regeneration of the dry deciduous forest of western Madagascar with the complete set of primary forest tree species seems to depend upon the presence of E. fulvus .     

Neighborhood and community interactions determine the spatial pattern of tropical tree seedling survival

Factors affecting survival and recruitment of 3531 individually mapped seedlings of Myristicaceae were examined over three years in a highly diverse neotropical rain forest, at spatial scales of 1-9 m and 25 ha. We found convincing evidence of a community compensatory trend (CCT) in seedling survival (i.e., more abundant species had higher seedling mortality at the 25-ha scale), which suggests that density-dependent mortality may contribute to the spatial dynamics of seedling recruitment. Unlike previous studies, we demonstrate that the CCT was not caused by differences in microhabitat preferences or life history strategy among the study species. In local neighborhood analyses, the spatial autocorrelation of seedling survival was important at small spatial scales (1-5 m) but decayed rapidly with increasing distance. Relative seedling height had the greatest effect on seedling survival. Conspecific seedling density had a more negative effect on survival than heterospecific seedling density and was stronger and extended farther in rare species than in common species. Taken together, the CCT and neighborhood analyses suggest that seedling mortality is coupled more strongly to the landscape-scale abundance of conspecific large trees in common species and the local density of conspecific seedlings in rare species. We conclude that negative density dependence could promote species coexistence in this rain forest community but that the scale dependence of interactions differs between rare and common species.     

Mortality of large trees and lianas following experimental drought in an Amazon forest

Severe drought episodes such as those associated with El Ni?o Southern Oscillation (ENSO) events influence large areas of tropical forest and may become more frequent in the future. One of the most important forest responses to severe drought is tree mortality, which alters forest structure, composition, carbon content, and flammability, and which varies widely. This study tests the hypothesis that tree mortality increases abruptly during drought episodes when plant-available soil water (PAW) declines below a critical minimum threshold. It also examines the effect of tree size, plant life form (palm, liana, tree) and potential canopy position (understory, midcanopy, overstory) on drought-induced plant mortality. A severe, four-year drought episode was simulated by excluding 60% of incoming throughfall during each wet season using plastic panels installed in the understory of a 1-ha forest treatment plot, while a 1-ha control plot received normal rainfall. After 3.2 years, the treatment resulted in a 38% increase in mortality rates across all stems >2 cm dbh. Mortality rates increased 4.5-fold among large trees (>30 cm dbh) and twofold among medium trees (10-30 cm dbh) in response to the treatment, whereas the smallest stems were less responsive. Recruitment rates did not compensate for the elevated mortality of larger-diameter stems in the treatment plot. Overall, lianas proved more susceptible to drought-induced mortality than trees or palms, and potential overstory tree species were more vulnerable than midcanopy and understory species. Large stems contributed to 90% of the pretreatment live aboveground biomass in both plots. Large-tree mortality resulting from the treatment generated 3.4 times more dead biomass than the control plot. The dramatic mortality response suggests significant, adverse impacts on the global carbon cycle if climatic changes follow current trends.     

Bat Use of Remnant Old-Growth Redwood Stands

Most of the old-growth redwood ( Sequoia sempervirens ) in California has been cut; regenerating forests will probably never resemble those that were harvested, and what old growth remains on private land occurs in small, isolated remnant patches. The landscapes in which these stands occur differ so markedly from their original condition that their value as habitat to many species of wildlife, including bats, is unknown. Previous research in unfragmented redwood forests demonstrated that bats use basal hollows in old-growth redwoods as roosts. We sought to determine whether bats use similar old-growth trees as roosts when they occur in small, remnant patches of isolated old growth on commercial forest land. We compared bat occurrence in remnant and contiguous stands by collecting guano in traps suspended in hollows and by monitoring flight activity with ultrasonic bat detectors. Hollows in trees within the remnant stands had significantly more guano deposited per tree than the trees within the contiguous forest. The mean numbers of bat passes per night were statistically indistinguishable between the two treatments, although mean flight activity in the remnant stands was greater than in the contiguous forest. Bats frequently used basal hollows in small (<5 ha) stands of remnant old growth, which may be due to the closer proximity of remnants to stream courses, to their greater interface with edge where foraging success may be greater, or to the fact that the lower density of hollow-bearing trees in remnants than in contiguous forest favored greater use per tree. Significant use of small, residual old-growth redwood provides reason to maintain these remnants in managed landscapes as potentially important habitat for forest bats.     

Potential site productivity influences the rate of forest structural development

Development and maintenance of structurally complex forests in landscapes formerly managed for timber production is an increasingly common management objective. It has been postulated that the rate of forest structural development increases with site productivity. We tested this hypothesis for Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) forests using a network of permanent study plots established following complete timber harvest of the original old-growth forests. Forest structural development was assessed by comparing empirical measures of live tree structure to published values for Douglas-fir forests spanning a range of ages and structural conditions. The rate of forest structural development--resilience--exhibited a positive relationship with site index, a measure of potential site productivity. Density of shade-intolerant conifers declined in all study stands from an initial range of 336-4068 trees/ha to a range of 168-642 trees/ha at the most recent measurement. Angiosperm tree species declined from an initial range of 40-371 trees/ha to zero in seven of the nine plots in which they were present. Trends in shade-tolerant tree density were complex: density ranged from 0 to 575 trees/ha at the first measurement and was still highly variable (25-389 trees/ha) at the most recent measurement. Multivariate analysis identified the abundance of hardwood tree species as the strongest compositional trend apparent over the study period. However, structural variables showed a strong positive association with increasing shade-tolerant basal area and little or no association with abundance of hardwood species. Thus, while tree species succession and forest structural development occur contemporaneously, they are not equivalent processes, and their respective rates are not necessarily linearly related. The results of this study support the idea that silvicultural treatments to accelerate forest structural development should be concentrated on lower productivity sites when the management objective is reserve-wide coverage of structurally complex forests. Alternatively, high-productivity sites should be prioritized for restoration treatments when the management objective is to develop structurally complex forests on a portion of the landscape.     

Conservation of Epiphyte Diversity in an Andean Landscape Transformed by Human Land Use

Abstract: Epiphytes are diverse and important elements of tropical forests, but as canopy‐dwelling organisms, they are highly vulnerable to deforestation. To assess the effect of deforestation on epiphyte diversity and the potential for epiphyte conservation in anthropogenically transformed habitats, we surveyed the epiphytic vegetation of an Ecuadorian cloud forest reserve and its surroundings. Our study was located on the western slopes of the Andes, a global center of biodiversity. We sampled vascular epiphytes of 110 study plots in a continuous primary forest; 14 primary forest fragments; isolated remnant trees in young, middle‐aged, and old pastures; and young and old secondary forests. It is the first study to include all relevant types of habitat transformation at a single study site and to compare epiphyte diversity at different temporal stages of fragmentation. Epiphyte diversity was highest in continuous primary forest, followed by forest fragments and isolated remnant trees, and lowest in young secondary forests. Spatial parameters of habitat transformation, such as fragment area, distance to the continuous primary forest, or distance to the forest edge from inside the forest, had no significant effect on epiphyte diversity. Hence, the influence of dispersal limitations appeared to be negligible or appeared to operate only over very short distances, whereas microclimatic edge effects acted only in the case of completely isolated trees, but not in larger forest fragments. Epiphyte diversity increased considerably with age of secondary forests, but species assemblages on isolated remnant trees were impoverished distinctly with time since isolation. Thus, isolated trees may serve for recolonization of secondary forests, but only for a relatively short time. We therefore suggest that the conservation of even small patches of primary forest within agricultural landscape matrices is essential for the long‐term maintenance of the high epiphyte diversity in tropical cloud forests.     

Effects of Habitat Fragmentation on Bird Species in a Relict Temperate Forest in Semiarid Chile

Abstract: We analyzed the structure and composition of a bird assemblage in a fragmented relict temperate forest located in northcentral Chile ( Fray Jorge National Park). In terms of species composition, the bird assemblage we found in Fray Jorge was more similar to southern temperate forest sites, located more than 1200 km south of Fray Jorge, than to localities found in nearby scrub habitats. The relict character and long-term isolation of the Fray Jorge forest provides a natural experiment with which to establish the potential long-term effects of fragmentation and isolation on southern Chilean temperate forests. Between May 1996 and March 1997, we conducted seasonal surveys of birds in six forest fragments, ranging in size from 0.5 to 22.5 ha, at Fray Jorge. The number of bird species at each forest fragment was positively correlated with fragment area during all seasons. The relict forest system had a steeper species-area slope than that reported for similar temperate-forest bird assemblages in forest fragments within Chiloé Island and for islands across the Chiloé Archipelago in southern Chile. In this regard, this bird fauna resembled a depauperate oceanic archipelago. This difference in area effects is likely a consequence of the minimization of rescue effects because of the absence of large source forest areas nearby and the long-term isolation of the system. In addition, the distribution of species among forest fragments in Fray Jorge was not random, showing a nested subset pattern. Thus, some species occur across all fragments, regardless of their area, and therefore are less affected by habitat fragmentation and less prone to local extinction. These results suggest that, for south-temperate forest birds, large fragments (or reserves) should afford better protection against extinction than small forest patches.     

Using Distance from Putative Source Woodlots to Predict Occurrence of Forest Birds in Putative Sinks

Abstract:  For some species of forest-breeding birds, productivity is much higher in large forest fragments than in small forest fragments and is higher than the level needed to replace individuals within the fragment (sources). Thus large forest fragments potentially provide excess individuals to neighboring small fragments where productivity is much lower and not adequate for replacement (sinks). We used occurrence data and distances between putative sources and sinks for four species of forest birds with this pattern of demography in large and small forest fragments to predict occupancy in small fragments. For the Ovenbird (  Seiurus aurocapilla ), Wood Thrush (  Hylocichla mustelina ), Veery ( Catharus fuscescens ), and Rose-breasted Grosbeak ( Pheucticus ludovicianus ), distance from large woodlots was a significant predictor of occurrence in small woodlots. Distance from large woodlots did not predict occupancy of Red-eyed Vireos (  Vireo olivaceus ), a species with no apparent difference in productivity in relation to fragment size. Neither vegetation features nor area of the small woodlots adequately explained patterns of occupancy for any species. These results suggest that maintenance of large, productive forest fragments benefit regional occurrence of forest-breeding birds by providing individuals to neighboring small woodlots. Limited reproduction in these small woodlots means that they may act as a reservoir for excess individuals in the case of catastrophic events that result in vacancies in source woodlots. Conservation and sound forest management of small forests should be directed particularly toward those closer to large regional forests.     

Influence of Forest Fragmentation on Community Structure of Frogs and Lizards in Northeastern Costa Rica

Abstract:  To better understand responses of reptiles and amphibians to forest fragmentation in the lowland Neotropics, we examined community and population structure of frogs and lizards in the fragmented landscape surrounding La Selva Biological Station in the Sarapiquí region of northeastern Costa Rica. We used diurnal quadrats and nocturnal transects to sample frogs and lizards in nine forest fragments (1–7 ha each) and La Selva (1100 ha). Species richness in all fragments combined was 85% of that found in La Selva with comparable sampling effort. Richness varied from 10 to 24 species among forest fragments, compared with 36 species at La Selva. Lizard density was higher and frog density was lower in forest fragments than in La Selva. Community composition varied among sites and by fragment size class, and species occurrence was nested with respect to fragment area. Isolation and habitat variables did not significantly affect species richness, composition, or nestedness. We classified 34% of species as fragmentation sensitive because they were absent or occurred at low densities in fragments. Nevertheless, the relatively high diversity observed in the entire set of fragments indicates that preserving a network of small forest patches may be of considerable conservation value to the amphibians and reptiles of this region.     

Complementary Roles of Home Gardens and Exotic Tree Plantations as Alternative Habitats for Plants of the Ethiopian Montane Rainforest

Abstract:  Many tropical forests have been converted for agri- or silviculture or a combination of both (agroforestry). Conservation at a landscape scale requires an understanding of the distribution and abundance of native biodiversity in these converted natural ecosystems, of which the knowledge is especially poor for African agroecosystems. We compared species density and species composition of four plant groups (trees and shrubs, epiphytic vascular plants, mosses, and liverworts) among three arboreal land-use types in southwestern Ethiopia (montane rainforest fragments, shade-tree coffee home gardens, and exotic tree plantations). Species density was significantly higher in forests than in coffee home gardens for all plant groups and in exotic tree plantations for all groups except mosses. Home gardens had more vascular epiphytic species than plantations, whereas the reverse was true for mosses and liverworts. The species composition of the forest plots was sometimes more similar to home-garden plots than plantation plots and sometimes vice versa. Fifteen forest plots had, however, cumulatively more species than a random selection of 15 nonforest (coffee home garden and plantation) plots, even if the 2 plot types complemented each other in terms of habitats for forest plants. Tree plantations dominated by Eucalyptus had many small trees and shrubs in common with forests, whereas plantations with Cupressus were important substrates for forests mosses and liverworts. Our results illustrate the importance of undisturbed forests habitats for conservation of species at a landscape scale and that different human-made land-use types may complement each other in their capacity as additional habitats for forest species.     

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     

Evaluation of sampling methods for the estimation of structural indices in forest stands

The paper is about the accurate (i.e. unbiased and precise) and efficient estimation of structural indices in forest stands. We present SIAFOR, a computer programme for the calculation of four nearest-neighbour indices, which describe the spatial arrangement of tree positions, the distribution pattern of species, and the size differentiation between trees. The study uses SIAFOR as a sampling simulator in eight completely stem-mapped forest stands of varying area and structural complexity. We statistically evaluate two sample types (distance and plot sampling), comparing sampling error, bias and minimum sample size for index estimation. We introduce the concepts of measurement expansion factor (MEF) and design expansion factor (DEF) for the technical evaluation of sample type efficiency (optimal sample type). Results indicate that sampling error can reach high levels and that minimum sample sizes for index estimation often amply exceed the limit of 20% of tree density or 20 trees per species per hectare, that we set as the highest feasible sample size in normal situations. We found clear feasibility limits (in terms of minimal tree densities and reachable accuracy levels) for the estimation of all investigated indices. Generally, equal or higher sample sizes are needed for plot sampling than for distance sampling to reach equal accuracy levels. Nevertheless, plot sampling resulted more efficient for the estimation of tree size differentiation at low to medium accuracy levels. For all other investigated indices distance sampling resulted more efficient than plot sampling. Minimum sample size increases with accuracy and is negatively correlated with tree density. At a given accuracy level minimum sample size is highest for the estimation of relative mingling and lowest for tree size differentiation; furthermore it is generally lower in large stands than in small ones. Because of the consistency of our conclusions in all of the investigated stands, we think they apply in most stands of similar area (between 1 and 10 ha) and species diversity (not more than four species).