Independent effects of habitat loss,habitat fragmentation and structural connectivity on forest‐dependent birds

Aim Habitat loss and fragmentation are amongst the greatest threats to biodiversity world‐wide. However, there is still little evidence on the relative influence of these two distinct processes on biodiversity, and no study, to date, has investigated the independent contribution of structural connectivity in addition to habitat loss and fragmentation. The aim of this study is to evaluate the independent effects of habitat loss (the decrease in total amount of habitat), habitat fragmentation per se (habitat subdivision) and structural connectivity (in the form of hedgerow networks) on the distribution of seven resident forest‐dependent birds in central Italy. Location Central Italy. Methods We strategically selected 30 landscapes (each of 16 km² in size) with decreasing total amount of forest cover and with contrasting configuration of patches and contrasting lengths of hedgerow networks. Presence/absence of birds in each landscape unit was studied through point counts. Results The amount of forest cover in the landscape had the strongest relative influence on birds’ occupancy, whilst habitat subdivision played a negligible role. Structural connectivity and the geographic position of the landscape unit played a relatively important role for four species. Main conclusions Our study shows the importance of disentangling the contribution of different landscape properties in determining distribution patterns. Our results are consistent with the fact that halting habitat loss and carrying out habitat restoration should be conservation priorities, since habitat loss is the main factor affecting the distribution of the target species; implementation of structural connectivity through hedgerows, instead, should be evaluated with caution since its contribution is secondary to the predominant role of habitat loss.     

A global assessment of primate responses to landscape structure

Land‐use change modifies the spatial structure of terrestrial landscapes, potentially shaping the distribution, abundance and diversity of remaining species assemblages. Non‐human primates can be particularly vulnerable to landscape disturbances, but our understanding of this topic is far from complete. Here we reviewed all available studies on primates' responses to landscape structure. We found 34 studies of 71 primate species (24 genera and 10 families) that used a landscape approach. Most studies (82%) were from Neotropical forests, with howler monkeys being the most frequently studied taxon (56% of studies). All studies but one used a site‐landscape or a patch‐landscape study design, and frequently (34% of studies) measured landscape variables within a given radius from the edge of focal patches. Altogether, the 34 studies reported 188 responses to 17 landscape‐scale metrics. However, the majority of the studies (62%) quantified landscape predictors within a single spatial scale, potentially missing significant primate–landscape responses. To assess such responses accurately, landscape metrics need to be measured at the optimal scale, i.e. the spatial extent at which the primate–landscape relationship is strongest (so‐called ‘scale of effect’). Only 21% of studies calculated the scale of effect through multiscale approaches. Interestingly, the vast majority of studies that do not assess the scale of effect mainly reported null effects of landscape structure on primates, while most of the studies based on optimal scales found significant responses. These significant responses were primarily to landscape composition variables rather than landscape configuration variables. In particular, primates generally show positive responses to increasing forest cover, landscape quality indices and matrix permeability. By contrast, primates show weak responses to landscape configuration. In addition, half of the studies showing significant responses to landscape configuration metrics did not control for the effect of forest cover. As configuration metrics are often correlated with forest cover, this means that documented configuration effects may simply be driven by landscape‐scale forest loss. Our findings suggest that forest loss (not fragmentation) is a major threat to primates, and thus, preventing deforestation (e.g. through creation of reserves) and increasing forest cover through restoration is critically needed to mitigate the impact of land‐use change on our closest relatives. Increasing matrix functionality can also be critical, for instance by promoting anthropogenic land covers that are similar to primates' habitat.     

Avian fitness consequences match habitat selection at the nest‐site and landscape scale in agriculturally fragmented landscapes

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Influence of within‐patch habitat quality on high‐Andean Polylepis bird abundance

Habitat restoration strategies for fragmented high Andean forest landscapes must consider the influence of within‐patch habitat quality on bird abundance. I examined vegetation and bird abundance at three locations within a highly fragmented Polylepis forest landscape in the Cordillera Vilcanota, southern Peru. Across the landscape, there was significant variation in the vegetation structure of Polylepis forest patches of different size categories, especially in terms of tree girth, tree height, tree density, and canopy vegetation structure. Principal Component Analysis extracted five factors of habitat quality, which together accounted for 74.2% of the variability within 15 habitat variables. Polylepis bird species differed in their responses to habitat quality but, overall, variation in Polylepis bird abundance was not fully captured by the range of habitat quality variables. Tall, dense vegetation cover was clearly important for 11 conservation‐important species, a high density of large trees was important for 10 species and primary forest ground cover was important for eight species. Habitat quality exhibited no significant influence on the abundance of only one species –Asthenes urubambensis. The abundance of seven species was associated with lower elevation forest, but only one species was associated with higher elevation forest. Management of habitat quality in large and medium remnant forest patches throughout the Cordillera Vilcanota, particularly in the 3800–4200 m elevation range, will be a cornerstone in ensuring the persistence of the majority of conservation‐important bird species populations.     

Impact of landscape spatial pattern on liana communities in tropical rainforests at Los Tuxtlas,Mexico

Questions: What are the species composition and species and stem densities of liana communities in tropical landscapes of different deforestation levels? Which spatial attributes (forest cover, patch area, shape and isolation) have the strongest influence on liana communities in these landscapes? Location: Forty‐five rainforest patches in Los Tuxtlas Biosphere Reserve, Mexico. Methods: In three landscapes with different deforestation levels (HDL=4%; IDL=11%; and LDL=24% of remaining forest cover) liana communities (DBH ≥2.5 cm) were characterized in 15 randomly selected patches per landscape (10 50 m × 2 m transects per patch=0.1 ha), and evaluated the effects of patch area, shape and isolation on liana species and stem density (number of species and stems per 0.1 ha). Results: A total of 64 taxa and 24 families were sampled. Species composition differed highly among landscapes, with HDL being the most dissimilar landscape. The response of lianas to landscape spatial pattern differed significantly among landscapes. Proximity to villages had a strong positive effect on species and stem densities in LDL and IDL. There was a sharp decrease in liana stem density in HDL, with four patches (27%) found to be unoccupied by lianas. Conclusions: Fragmentation may have a positive effect on lianas, partly because of edge effects. This positive effect seems to be limited by the proportion of remaining forest cover in the landscape, as the liana communities had collapsed in the most deforested landscape.     

Biodiversity and ecological forest-cover domains in boreal landscapes

Studies of spatial forest patterns have traditionally focused on habitat remnants within a landscape, but few have explicitly accounted for natural habitat patterns inherent to those landscapes. At broader scales, all cover types are to some degree subdivided. In boreal forest landscapes, forests, peat lands and water bodies form a variety of different spatial structures depending on their proportions and arrangement. To assess the spatial arrangement of forest patches relevant to organisms and ecological processes, we systematically sampled and analysed 57 boreal forest landscapes (10,000 ha in size) in central Finland. Our results show that even though forest is spatially very subdivided into discrete patches in boreal landscapes it becomes generally well-connected if narrow non-forested gaps are ignored. The proportion of forest cover varied from 17.8 to 75.3% and the number of discrete forest patches from 37 to 213. The average percolation threshold for forest cover was 46.8%. If ≤100 m wide non-forested gaps were ignored and forest patches were joined, the percolation threshold dropped to 33.3%. There were on average 13 discrete clusters of forest patches if forest patches within 200 m distance were combined. These results suggest that if a boreal forest species is able to cross even relatively narrow non-forested gaps, it is likely to perceive these boreal forest landscapes as continuous. Even though our present analysis was based on static forest cover patterns, it is important to consider landscape pattern domains when assessing habitat fragmentation and its consequences to populations of organisms.     

Landscape mosaic of Araucaria forest and forest monocultures influencing understorey spider assemblages in southern Brazil

Abstract This study investigates how abundance, diversity and composition of understorey spiders were influenced by four different forest habitats in a southern Brazilian Araucaria forest. The study area encompasses a landscape mosaic comprised of Araucaria forest, Araucaria plantation, Pinus plantation, and Eucalyptus plantation. Understorey spiders were collected by beating the vegetation inside three patches of each forest habitat. To assess possible predictors of spider assemblage structure, several patch features were analysed: potential prey abundances, estimation of vegetation cover, diversity index of vegetation types, patch ages, patch areas, and geographical distance between patches. To assess the influence of high‐level taxa approaches on spider assemblage patterns, analyses were carried out individually for family, genera and species levels. Additionally, Mantel tests were carried out in underlying similarity matrices between each taxon. Significant differences in spider abundances among forest habitats were found. Pinus plantations showed the highest abundance of spiders and Eucalyptus plantations showed the lowest abundance. Spider abundance was significantly influenced by patch ages, geographical distance and vegetation cover. Expected numbers of families, genera and species did not vary among forest habitats. Spider composition of two Eucalyptus patches differed from the other forest patches, probably due to their low vegetation cover and isolation. Genera composition was the best correlate of species composition, showing that a higher‐level surrogate can be an alternative to the species approach. The understorey spider diversity in this managed area could be maintained when suitable habitat structures are provided, thus ensuring the connectivity between different habitat types. Further studies should focus on individual species responses to the conversion of native forest to monocultures.     

The effect of landscape pattern on Mediterranean vegetation dynamics: A modelling approach using functional types

Abstract. In the framework of land use changes in the Mediterranean area, I asked to what extent different landscape structures might determine long‐term dynamics in Mediterranean ecosystems. To answer this question, a spatially explicit model was developed (the Melca model), incorporating two functional types of woody species dominant in Mediterranean ecosystems: a resprouter (R) and a non‐resprouter fire‐recruiter (seeders, S). The model was used as a tool for generating hypotheses on the possible consequences of different landscape scenarios. Thus, five different hierarchically structured random landscapes were generated, all having the same cover for the two functional types but different landscape structure (ranging from highly heterogeneous to homogeneous landscapes). After a 100‐yr simulation, plant cover and spatial pattern had changed and the changes were different for the different initial spatial configurations, suggesting that long‐term vegetation dynamics is spatially dependent (the resultant dynamics are sensitive to the initial spatial structure). In the landscapes where R‐type species had a low number of large patches and S‐species had a large number of small patches, the number of R‐patches increased and their size decreased, while the number of S‐patches decreased. In these cases, the final cover of the two types changed little from the initial cover. Landscapes with a large number of small R‐patches interspersed with S‐patches had a decrease in the number of R‐patches, an increase in the number of S‐patches and a decrease in the size of S‐patches. In these landscapes, final cover was significantly changed, increasing in R‐type and decreasing in S‐type species. These results suggest that low spatial autocorrelation (low aggregation) favours R‐type species. Implications for land management are also discussed.     

Isolation determines patterns of species presence in highly fragmented landscapes

In fragmented landscapes, changes in habitat availability, patch size, shape and isolation may affect survival of local populations. Proposing efficient conservation strategies for such species relies initially on distinguishing the particular effects of those factors. To address these issues, we investigated the occurrence of 3 bird species in fragmented Brazilian Atlantic Forest landscapes. Playback techniques were used to collect presence/absence data of these species inside 80 forest patches, and incidence models were used to infer their occupancy pattern from landscape spatial structure. The relative importance of patch size, shape and surrounding forest cover and isolation was assessed using a model selection approach based on maximum likelihood estimation. The presence of all species was in general positively affected by the amount of surrounding habitat and negatively affected by inter‐patch distances. The joint effects of patch size and the surrounding landscape characteristics were important determinants of occupancy for two species. The third species was affected only by forest cover and mean patch isolation. Our results suggest that local species presence is in general more influenced by the isolation from surrounding forests than by patch size alone. We found evidence that, in highly fragmented landscapes, birds that can not find patches large enough to settle may be able to overcome short distances through the matrix and include several nearby patches within their home‐ranges to complement their resource needs. In these cases, patches must be defined as functionally connected habitat networks rather than mere continuous forest segments. Bird conservation strategies in the Atlantic forest should focus on increasing patch density and connectivity, in order to implement forest networks that reduce the functional isolation between large remnants with remaining core habitat.     

Chainsawing for conservation: Ecologically informed tree removal for habitat management

Summary In many ecosystems, increases in vegetation density and the resulting closure of forest canopies are threatening the viability of species that depend upon open, sunlight‐exposed habitats. Consequently, we need to develop management strategies that recreate open habitats while minimizing the impacts on non‐target areas. Selective logging creates canopy gaps, but may result in undesirable effects in other respects. Thus, chainsaws have not been a popular tool for conservation. We conducted a landscape‐scale experiment to test whether selective tree removal can restore patch‐level habitat quality for Australia’s most endangered snake (Hoplocephalus bungaroides) and its main prey (the lizard Oedura lesueurii). We selectively removed canopy trees surrounding 25 overgrown rock outcrops and compared the resultant habitat structure and abiotic conditions to 30 overgrown, shady outcrops and 20 open, sunny outcrops. Removing vegetation decreased canopy cover by 19% in experimental plots and increased incident radiation and thermal regimes. These changes increased the availability of suitable shelter sites for our target species by 131%. At the landscape scale, our manipulations had a trivial effect on forest habitat; by increasing the area of sun‐exposed outcrops, we decreased forest cover by <0.1%. Our results show that targeted canopy removal can increase the availability of sun‐exposed habitat patches for endangered species in biologically meaningful ways. Thus, selective tree felling may be an effective conservation tool for open‐habitat specialists threatened by vegetation overgrowth.     

Agricultural intensification alters bat assemblage composition and abundance in a dynamic Neotropical landscape

The recent trend of agricultural intensification in tropical landscapes poses a new threat to biodiversity conservation. Conversion of previously heterogeneous agricultural landscapes to intensive plantation agriculture simplifies and homogenizes the landscape, reducing availability, and connectivity of natural habitat for native species. To assess the impact of agricultural intensification on bats, we characterized the bat assemblage in the Sarapiquí region of Costa Rica, where heterogeneous land uses are being converted to intensive, large‐scale pineapple plantations. In 2012 and 2013, we sampled bats in 20 remnant forest patches surrounded by varying proportions of pasture, mature forest, and pineapple and captured 1821 individual bats representing 39 species. We used ordination analyses to evaluate changes in species composition, where pineapple is the main component of the agricultural matrix. We identified landscape metrics specifically correlated with pineapple and used multiple linear regression to test their effects on bat species richness, diversity, and guild‐specific relative abundance. Results suggest pineapple expansion is driving changes in assemblage composition in remnant forest patches, resulting in new assemblages with higher proportions of frugivorous bats and lower proportions of insectivorous bats than in continuous mature forests. In addition, while pineapple does not diminish total bat species richness and diversity, the reduced forest cover and increased distance between forest patches in pineapple plantations has a significant negative impact on the relative abundance of insectivores. We also identify a potential threshold effect whereby patches surrounded by more than 50 percent forest can retain assemblage composition similar to that found in continuous mature forest.     

Additive partitioning of butterfly diversity in a fragmented landscape: importance of scale and implications for conservation

Aim Most of the Atlantic Forest in Brazil occurs in fragments of various sizes. Previous studies indicate that forest fragmentation affects fruit‐feeding butterflies. Conservation strategies that seek to preserve organisms that are distributed in high‐fragmented biomes need to understand the spatial distribution of these organisms across the landscape. In view of the importance of understanding the fauna of these forest remnants, the objective of the present work is to investigate the extent to which the diversity of this group varies across spatial scales ranging from within‐forest patches to between landscapes. Location South America, south‐eastern Brazil, São Paulo State. Methods We used bait traps to sample fruit feeding butterflies at 50 points in 10 fragments in two different landscapes during a period of 12 months. Total species richness and Shannon index were partitioned additively in diversity at trap level, and beta diversity was calculated among traps, among forest patches, and between landscapes. We used permutation tests to compare these values to the expected ones under the null hypothesis that beta diversity is only a random sampling effect. Results There was significant beta diversity at the smallest scale examined; however, the significance at higher scales depends on the diversity measurement used. Beta diversity with Shannon index was smaller than expected by chance among fragments, whereas species richness was not. Among landscapes, only beta diversity in richness was higher than expected by chance. Main conclusions The results observed occur because there is great variability in species composition among forest patches in the same landscape, changing this diversity even though the communities are formed from the same pool of species. At the largest scale evaluated (between landscapes), these pattern changes and differences in beta diversity in richness were detectable. This difference is probably caused by the presence of rare species. Thus, a conservation strategy that seeks to preserve as many species as possible per unit of area in high‐fragmented biomes should give priority to protecting fragments in different landscapes, rather than more fragments in the same landscape.     

Permeability of Neotropical agricultural lands to a key native ungulate—Are well‐connected forests important?

Much of what remains of the Earth's tropical forests is embedded within agricultural landscapes, where forest is reduced and fragmented. As native forest ungulates are critical to maintaining forest function, it is imperative to understand how this functional group responds to declines in forest cover and connectivity resulting from agricultural expansion. We addressed this issue by evaluating selection of forest cover and forest connectivity by a key native ungulate of Neotropical forests, the white‐lipped peccary (Tayassu pecari Link 1795, Tayassuidae, Cetartiodactyla), in agricultural landscapes of Brazil. We evaluated selection using compositional analysis at two hierarchical levels, landscape, and home range. From 2013 to 2019, we GPS‐tracked eight white‐lipped peccary herds in Southwest Brazil, resulting in a total of 14,460 GPS locations. We found that herds can live in landscapes with a wide range of forest cover (35%–81% of home ranges covered by native forest), with significant, but not strong, selection at the landscape level (p = .045). Nevertheless, herds strongly select for forest cover within their home ranges (81%–97% of locations within native forest; highly significant selection at the home‐range level: p = .008). As for connectivity, herds significantly select the largest, most connected forest fragments at the landscape level (p = .04), but not at the home‐range level (p = .07). Our results support that Neotropical forests within agricultural landscapes need to be well connected in order to preserve this key native ungulate and maintain long‐term forest function. Abstract in Portuguese is available with online material.     

The roles of landscape context, niche breadth, and range boundaries in predicting species responses to habitat alteration

Extant species in human‐dominated landscapes differ in their sensitivity to habitat loss and fragmentation, although extinctions induced by environmental alteration reduce variation and result in a surviving subset of species with some degree of ‘resistance’. Here, we test the degree to which variable responses to habitat alteration are (1) essentially an inherent property of a taxon subject to constraints imposed by its geographical range, as suggested by Swihart et al. (2003), (2) a function of the landscape in which a species occurs, or (3) a function of spatial trends occurring on large scales. We used data collected on 33 vertebrate species during 2001–04 across the upper Wabash River basin, Indiana, in 35 square ‘landscapes’, each 23 km² in size. Six species of forest rodent, six species of grassland rodents, seven species of bats, eight species of aquatic turtles, and six species of amphibians were sampled at 504, 212, 590, 228, and 625 patches, respectively. The fraction of patches of primary habitat (e.g. forests for tree squirrels, wetlands for aquatic turtles) occupied by a target species was used as a response variable. On a basin‐wide scale, 47% of variation in proportional occupancy among species could be explained by taxon‐specific variables; occupancy rates were related positively to niche breadth and negatively to the proximity of a geographical range boundary. After controlling for species effects, landscape‐level occupancy rates varied significantly for 16 of 33 species, with variation partitioned among landscape variables alone (mean = 11% of variation), spatial trend variables alone (26%), and both variable sets jointly (8%). Among landscape variables, percentage forest cover positively affected occupancy rates of three bat species and a tree squirrel. Variation in occupancy rates among landscapes was consistent with large‐scale spatial trends for 13 species. Our findings demonstrate the general importance of niche breadth as a predictor of species responses to habitat alteration and highlight the importance of viewing the effects of habitat loss and fragmentation at multiple spatial scales.     

Genetic evidence for landscape effects on dispersal in the army ant Eciton burchellii

Inhibited dispersal, leading to reduced gene flow, threatens populations with inbreeding depression and local extinction. Fragmentation may be especially detrimental to social insects because inhibited gene flow has important consequences for cooperation and competition within and among colonies. Army ants have winged males and permanently wingless queens; these traits imply male‐biased dispersal. However, army ant colonies are obligately nomadic and have the potential to traverse landscapes. Eciton burchellii, the most regularly nomadic army ant, is a forest interior species: colony raiding activities are limited in the absence of forest cover. To examine whether nomadism and landscape (forest clearing and elevation) affect population genetic structure in a montane E. burchellii population, we reconstructed queen and male genotypes from 25 colonies at seven polymorphic microsatellite loci. Pairwise genetic distances among individuals were compared to pairwise geographical and resistance distances using regressions with permutations, partial Mantel tests and random forests analyses. Although there was no significant spatial genetic structure in queens or males in montane forest, dispersal may be male‐biased. We found significant isolation by landscape resistance for queens based on land cover (forest clearing), but not on elevation. Summed colony emigrations over the lifetime of the queen may contribute to gene flow in this species and forest clearing impedes these movements and subsequent gene dispersal. Further forest cover removal may increasingly inhibit Eciton burchellii colony dispersal. We recommend maintaining habitat connectivity in tropical forests to promote population persistence for this keystone species.     

Quantitative changes of forest landscapes over the last century across Italy

A key topic in landscape ecology and vegetation science is the quantitative analysis of changes in forest cover over time, through the use of geomatics monitoring tools. Ecologists and landscape researchers are pointing out that a full understanding of ecosystems and landscapes should be based on the analysis of their functioning over long time series. Under this perspective, a long-term historical reconstruction of forest cover is essential. This study has aimed at examining the long-term dynamics of forest landscapes in Italy, over the last century, using recent remote-sensing based map (2012) and an accurate historical map (1936). A forest-non forest approach has been followed by the computation of a variety of landscape metrics using two analysis tools, with the final objective of quantifying changes in forest cover patterns and in the composition of specific landscape elements. Results show that forest landscape structure has significantly changed across Italy, resulting in a general trend of decreasing fragmentation and patchiness, mainly through enlargement of existing forest patches, which have also assumed a more geometrically regular shape. In relative terms, the greatest expansion of forest areas has occurred mainly in lowland districts characterised by the highest level of human pressure in the country.     

Densities of Bornean orang‐utans (Pongo pygmaeus morio) in heavily degraded forest and oil palm plantations in Sabah,Borneo

The conversion of forest to agriculture continues to contribute to the loss and fragmentation of remaining orang‐utan habitat. There are still few published estimates of orang‐utan densities in these heavily modified agricultural areas to inform range‐wide population assessments and conservation strategies. In addition, little is known about what landscape features promote orang‐utan habitat use. Using indirect nest count methods, we implemented surveys and estimated population densities of the Northeast Bornean orang‐utan (Pongo pygmaeus morio) across the continuous logged forest and forest remnants in a recently salvage‐logged area and oil palm plantations in Sabah, Malaysian Borneo. We then assessed the influence of landscape features and forest structural metrics obtained from LiDAR data on estimates of orang‐utan density. Recent salvage logging appeared to have a little short‐term effect on orang‐utan density (2.35 ind/km ²), which remained similar to recovering logged forest nearby (2.32 ind/km ²). Orang‐utans were also present in remnant forest patches in oil palm plantations, but at significantly lower numbers (0.82 ind/km ²) than nearby logged forest and salvage‐logged areas. Densities were strongly influenced by variation in canopy height but were not associated with other potential covariates. Our findings suggest that orang‐utans currently exist, at least in the short‐term, within human‐modified landscapes, providing that remnant forest patches remain. We urge greater recognition of the role that these degraded habitats can have in supporting orang‐utan populations, and that future range‐wide analyses and conservation strategies better incorporate data from human‐modified landscapes.     

Patch occupancy and abundance of local populations in landscapes differing in degree of habitat fragmentation: a case study of the colonial black‐headed gull,Chroicocephalus ridibundus

Aim This study investigated whether habitat fragmentation at the landscape level influences patch occupancy and abundance of the black‐headed gull, Chroicocephalus ridibundus, and whether the response of the species to environmental factors is consistent across replicated landscape plots. Location Water bodies (habitat patches) in southern Poland. Methods Surveys were conducted in two landscape types (four plots in each): (1) more‐fragmented landscape, in which habitat patches were small (mean size 2.2–6.2 ha) and far apart (mean distance 2.5–3.1 km); and (2) less‐fragmented landscape, in which habitat patches were large (mean size 9.2–16.5 ha) and separated by short distances (mean 0.9–1.4 km). Observations were performed twice in 284 potential habitat patches during the 2007 breeding season. Results Colonies were significantly more frequent and larger in the less‐fragmented landscapes than in the more‐fragmented ones. Probability of patch occupancy and number of breeding birds were positively related with patch size and these relationships were especially strong in the more‐fragmented landscapes. In the less‐fragmented landscapes, the occurrence of black‐headed gulls was negatively related to the distance to the nearest local population, but in the more‐fragmented landscapes such a relationship was not detected. As distance to the nearest habitat patch increased, the probability of the patch occupancy decreased in the more‐fragmented landscapes. Moreover, abundance was negatively influenced by distance to the nearest habitat patch, especially strongly in more‐fragmented landscapes. Proximity of corridors (rivers) positively influenced the occupation of patches regardless of landscape type. The number of islets positively influenced occupancy and abundance of local populations, and this relationship was stronger in the more‐fragmented landscapes. Main conclusions Our results are in agreement with predictions from metapopulation theory and are the first evidence that populations of black‐headed gulls may have a metapopulation structure. However, patch occupancy and abundance were differentially affected by explanatory variables in the more‐fragmented landscapes than in the less‐fragmented ones. This implies that it is impossible to derive, a priori, predictions about presence/abundance patterns based on only a single landscape.     

Immigration rates in fragmented landscapes--empirical evidence for the importance of habitat amount for species persistence

Background

The total amount of native vegetation is an important property of fragmented landscapes and is known to exert a strong influence on population and metapopulation dynamics. As the relationship between habitat loss and local patch and gap characteristics is strongly non-linear, theoretical models predict that immigration rates should decrease dramatically at low levels of remaining native vegetation cover, leading to patch-area effects and the existence of species extinction thresholds across fragmented landscapes with different proportions of remaining native vegetation. Although empirical patterns of species distribution and richness give support to these models, direct measurements of immigration rates across fragmented landscapes are still lacking.

Methodology/Principal Findings

Using the Brazilian Atlantic forest marsupial Gray Slender Mouse Opossum (Marmosops incanus) as a model species and estimating demographic parameters of populations in patches situated in three landscapes differing in the total amount of remaining forest, we tested the hypotheses that patch-area effects on population density are apparent only at intermediate levels of forest cover, and that immigration rates into forest patches are defined primarily by landscape context surrounding patches. As expected, we observed a positive patch-area effect on M. incanus density only within the landscape with intermediate forest cover. Density was independent of patch size in the most forested landscape and the species was absent from the most deforested landscape. Specifically, the mean estimated numbers of immigrants into small patches were lower in the landscape with intermediate forest cover compared to the most forested landscape.

Conclusions/Significance

Our results reveal the crucial importance of the total amount of remaining native vegetation for species persistence in fragmented landscapes, and specifically as to the role of variable immigration rates in providing the underlying mechanism that drives both patch-area effects and species extinction thresholds.     

Forest fragmentation and loss reduce richness,availability, and specialization in tropical hummingbird communities

Hummingbirds are important pollinators of many native Neotropical plants but their abundance and diversity in landscapes dominated by intensive human uses such as agriculture have rarely been examined, despite such land‐uses prevailing in the tropics. We examined how tropical deforestation affects hummingbird community structure in premontane forest patches embedded in a tropical countryside of Coto Brus Canton, Costa Rica. We captured hummingbirds in fourteen landscapes representing a gradient in patch size and forest amount, and tested for the effects of these variables on (1) hummingbird captures at flowers (pollinator availability); (2) species richness; and (3) filtering of functional traits. After accounting for sampling effects, both hummingbird availability and species richness declined by 40% and 50%, respectively, across the gradient in deforestation that we observed (9–66% forest within 1000 m). Focal patch size was the strongest predictor, even after statistically accounting for the amount of forest and matrix composition of landscapes. These reductions in availability and richness were well predicted by functional traits; morphologically specialized species with the capacity to transport long‐distance outcrossed pollen and low functional redundancy within the pollinator network showed the greatest sensitivity to landscape change. We hypothesize that declines in hummingbird availability, diversity, and functional traits are important mechanisms driving the observed pollen limitation of ornithophilous flowers in fragmented tropical landscapes. Efforts to conserve large forest patches and enhance matrix permeability are critical for maintaining forest hummingbird communities and pollination services under current and predicted deforestation regimes.