Beyond the Fragmentation Threshold Hypothesis: Regime Shifts in Biodiversity Across Fragmented Landscapes

Ecological systems are vulnerable to irreversible change when key system properties are pushed over thresholds, resulting in the loss of resilience and the precipitation of a regime shift. Perhaps the most important of such properties in human-modified landscapes is the total amount of remnant native vegetation. In a seminal study Andren proposed the existence of a fragmentation threshold in the total amount of remnant vegetation, below which landscape-scale connectivity is eroded and local species richness and abundance become dependent on patch size. Despite the fact that species patch-area effects have been a mainstay of conservation science there has yet to be a robust empirical evaluation of this hypothesis. Here we present and test a new conceptual model describing the mechanisms and consequences of biodiversity change in fragmented landscapes, identifying the fragmentation threshold as a first step in a positive feedback mechanism that has the capacity to impair ecological resilience, and drive a regime shift in biodiversity. The model considers that local extinction risk is defined by patch size, and immigration rates by landscape vegetation cover, and that the recovery from local species losses depends upon the landscape species pool. Using a unique dataset on the distribution of non-volant small mammals across replicate landscapes in the Atlantic forest of Brazil, we found strong evidence for our model predictions - that patch-area effects are evident only at intermediate levels of total forest cover, where landscape diversity is still high and opportunities for enhancing biodiversity through local management are greatest. Furthermore, high levels of forest loss can push native biota through an extinction filter, and result in the abrupt, landscape-wide loss of forest-specialist taxa, ecological resilience and management effectiveness. The proposed model links hitherto distinct theoretical approaches within a single framework, providing a powerful tool for analysing the potential effectiveness of management interventions.

Impact of seedling removal on regenerating community structure of a seasonal semideciduous forest

(Impact of seedling removal on regenerating community structure of a seasonal semideciduous forest). Transplanting seedlings and saplings from natural forests has been considered an alternative to producing saplings of native species for forest restoration purposes, but the possible impact of this procedure on plant community regeneration has not been investigated. This work evaluates the impact of different treatments of shrub and tree-seedling (up to 30 cm) removal from a seasonal semideciduous forest fragment located in southeastern Brazil on the natural regeneration process. Eighty 2x2 m plots were installed in two habitats (forest edge and interior) and submitted to four seedling-removal treatments (I, II - 100% removal with or without soil mixing; III - 50% removal without soil mixing: and IV - control treatment Without seedling removal). Regeneration density and richness were evaluated before treatment as well as 6, 12 and 18 months later. The results were compared among treatments for each evaluation period and among periods within treatments. There were similarities between edge and interior. The natural regeneration process did not improve with soil mixing. Plots submitted to seedling removal partially recovered plant density; however, these plots had lower species richness when compared to the control and to the initial values before treatment. Seedling removal has a negative impact on the regeneration process of low-density species, thus the use of natural regeneration as a sapling source for forest restoration purposes should focus only on high-density species with well-known regeneration strategies and not on the community as a whole.

Can intensive management accelerate the restoration of Brazil`s Atlantic forests?

Only 7% of the once extensive forest along the eastern coast of Brazil remains, and much of that is degraded and threatened by agricultural expansion and urbanization. We wondered if methods similar to those developed to establish fast-growing Eucalyptus plantations might also work to enhance survival and growth of rainforest species on degraded pastures composed of highly competitive C(4) grasses. An 8-factor experiment was laid out to contrast the value of different intensities of cultivation, application of fertilizer and weed control on the growth and survival of a mixture of 20 rainforest species planted at two densities: 3 m x 1 m, and 3 m x 2 m. Intensive management increased seedling survival from 90% to 98%, stemwood production and leaf area index (LAI) by similar to 4-fold, and stemwood production per unit of light absorbed by 30%. Annual growth in stem biomass was closely related to LAI alone (r(2) = 0.93, p < 0.0001), and the regression improved further in combination with canopy nitrogen content (r(2) =0.99, p < 0.0001). Intensive management resulted in a nearly closed forest canopy in less than 4 years, and offers a practical means to establish functional forests on abandoned agricultural land. (C) 2009 Elsevier B.V. All rights reserved.

Can native vegetation recover after slash pine cultivation in the Brazilian Savanna?

There is a widespread view that forest plantations with exotic species are green deserts, unable to sustain biodiversity. Few studies have demonstrated, however, that planted stands of exotic trees have a greater negative effect on the plant diversity of savanna vegetation. We compared the native woody flora under four stands of slash pine of about 45 years old with four stands where the previously existing native Cerrado vegetation was preserved and protected from disturbances for the same period, has changed into dense vegetation - the ""cerradao"", at Assis municipality, Sao Paulo State, Brazil. Aiming at understanding the potential ecological filters driving these communities, we assessed air and soil humidity, light availability and classified the native species on the basis of shade tolerance, dispersal syndrome and biomes in which they occur (Atlantic Forest or Cerrado). We recorded an average of 70 (+/- 13) species under pine stands and 54 (+/- 16) species in cerradao. Of the total of 136 species recorded, 78 occurred in both habitats, eight were exclusive to the ""cerradao"" (shade tolerant and also occurring in forest ecosystems) and 18 were recorded only under pine stands (82% heliophytic, exclusive to the Cerrado biome). Among the functional attributes and abiotic variables analyzed, only light availability explained the floristic differences found. Since richness was higher under pine, we refuted the hypothesis that exotic species constrain the establishment of the native species richness in the understory. On the other hand, the dark environment under the closed-canopy of the ""cerradao"" acts as a filter inhibiting the establishment of typical Cerrado species. Since pine stands, if managed in long cycle, maintain a reasonable pool of Cerrado endemic species in the understory pine plantations may be a good starting point for savanna restoration. (C) 2011 Elsevier B.V. All rights reserved.

Development and characterization of microsatellite markers for Psychotria tenuinervis (Rubiaceae), a shrub species from the Atlantic forest, and primers transferability from Coffea

The shrub species Psychotria tenuinervis (Rubiaceae) is native to the Brazilian Atlantic forest and is largely found within natural and disturbed forest fragments. Aiming to develop studies on population genetic structure of forest fragment species, eigth microsatellite markers were developed for P. tenuinervis. Also, 15 loci already developed for Coffea (Rubiaceae) were tested for transferability to this species. We utilized 45 individuals from natural populations of three different fragments-anthropic edge, interior fragment and natural edge, within the Brazilian Atlantic forest. The average number of alleles per locus was 2.5 (two-four alleles/locus). These loci will be useful for future population genetic studies aiming to the conservation and management of this species.

Temporal Diversity Patterns and Phenology in Fruit-feeding Butterflies in the Atlantic Forest

The Atlantic Forest deserves special attention due to its high level of species endemism and degree of threat. As in other tropical biomes, there is little information about the ecology of the organisms that occur there. The objectives of this study were to verify how fruit-feeding butterflies are distributed through time, and the relation with meteorological conditions. Species richness and Shannon index were partitioned additively at the monthly level, and beta diversity, used as a hierarchical measure of temporal species turnover, was calculated among months, trimesters, and semesters. Circular analysis was used to verify how butterflies are distributed along seasons and its relation with meteorological conditions. We sampled 6488 individuals of 73 species. Temporal diversity of butterflies was more grouped than expected by chance among the months of each trimester. Circular analyses revealed that diversity is concentrated in hot months (September-March), with the subfamily Brassolinae strongly concentrated in February-March. Average temperature was correlated with total abundance of butterflies, abundance of Biblidinae, Brassolinae and Morphinae, and richness of Satyrinae. The present results show that 3mo of sampling between September and March is enough to produce a nonbiased sample of the local assemblage of butterflies, containing at least 70 percent of the richness and 25 percent of abundance. The influence of temperature on sampling is probably related to butterfly physiology. Moreover, temperature affects resource availability for larvae and adults, which is higher in hot months. The difference in seasonality patterns among subfamilies is probably a consequence of different evolutionary pressures through time.

Prospects for biodiversity conservation in the Atlantic Forest: Lessons from aging human-modified landscapes

Recent global assessments have shown the limited coverage of protected areas across tropical biotas, fuelling a growing interest in the potential conservation services provided by anthropogenic landscapes. Here we examine the geographic distribution of biological diversity in the Atlantic Forest of South America, synthesize the most conspicuous forest biodiversity responses to human disturbances, propose further conservation initiatives for this biota, and offer a range of general insights into the prospects of forest species persistence in human-modified tropical forest landscapes worldwide. At the biome scale, the most extensive pre-Columbian habitats across the Atlantic Forest ranged across elevations below 800 masl, which still concentrate most areas within the major centers of species endemism. Unfortunately, up to 88% of the original forest habitat has been lost, mainly across these low to intermediate elevations, whereas protected areas are clearly skewed towards high elevations above 1200 masl. At the landscape scale, most remaining Atlantic Forest cover is embedded within dynamic agro-mosaics including elements such as small forest fragments, early-to-late secondary forest patches and exotic tree mono-cultures. In this sort of aging or long-term modified landscapes, habitat fragmentation appears to effectively drive edge-dominated portions of forest fragments towards an early-successional system, greatly limiting the long-term persistence of forest-obligate and forest-dependent species. However, the extent to which forest habitats approach early-successional systems, thereby threatening the bulk of the Atlantic Forest biodiversity, depends on both past and present landscape configuration. Many elements of human-modified landscapes (e.g. patches of early-secondary forests and tree mono-cultures) may offer excellent conservation opportunities, but they cannot replace the conservation value of protected areas and hitherto unprotected large patches of old-growth forests. Finally, the biodiversity conservation services provided by anthropogenic landscapes across Atlantic Forest and other tropical forest regions can be significantly augmented by coupling biodiversity corridor initiatives with biota-scale attempts to plug existing gaps in the representativeness of protected areas. (C) 2010 Elsevier Ltd. All rights reserved.

Are corridors, fragment size and forest structure important for the conservation of leaf-litter lizards in a fragmented landscape?

To investigate the implications of forest fragmentation for conservation of leaf-litter lizards the importance of fragment size, corridors and forest structure was examined in 20 forest fragments and six localities within a continuous forest in the Atlantic Plateau of Sao Paulo state, Brazil. The fragments were 2-276 ha in area and had different degrees of connectivity depending on the presence or absence of corridors. Two species of lizards were dominant, Ecpleopus gaudichaudii and Enyalius perditus. Variation in forest structure among sites was important only in explaining the abundance of E. perditus. Regardless of variation in forest structure, lizard species composition, total lizard abundance, number of species and abundance of E. perditus were sensitive to fragmentation per se but not to fragment size or corridor linkage. The inhospitable matrix surrounding fragments is probably what determines the presence and abundance of E. perditus and the high er lizard richness in continuous forests. These conditions may have prevented lizard species from recolonizing the forest fragments. Our results emphasize that the conservation of this leaf-litter fauna depends on the maintenance of large tracts of continuous forests and not on the size of fragments or on the presence of forest connections. Strategies for conservation of leaf-litter lizards in such highly fragmented Atlantic Forest landscapes should consider the enlargement of landscape connectivity between fragments and continuous forest, allowing the latter areas to act as a source of individuals for fragments.

Relative effects of fragment size and connectivity on bird community in the Atlantic Rain Forest: Implications for conservation

It is known that large fragment sizes and high connectivity levels are key components for maintaining species in fragments; however, their relative effects are poorly understood, especially in tropical areas. In order to test these effects, we built models for explaining understory birds occurrence in a fragmented Atlantic Rain Forest landscape with intermediate habitat cover (3%). Data from over 9000 mist-net hours from 17 fragments differing in size (2-175 ha) and connectivity (considering corridor linkages and distance to nearby fragments) were ranked under a model selection approach. A total 1293 individuals of 62 species were recorded. Species richness, abundance and compositional variation were mainly affected by connectivity indices that consider the capacity of species to use corridors and/or to cross short distances up to 30 m through the matrix. Bird functional groups were differently affected by area and connectivity: while terrestrial insectivores, omnivores and frugivores were affected by both area and connectivity, the other groups (understory insectivores, nectarivores, and others) were affected only by connectivity. In the studied landscape, well connected fragments can sustain an elevated number of species and individuals. Connectivity gives the opportunity for individuals to use multiple fragments, reducing the influence of fragment size. While preserving large fragments is a conservation target worldwide and should continue to be, our results indicated that connectivity between fragments can enhance the area functionally connected and is beneficial to all functional groups and therefore should be a conservation priority. (C) 2008 Elsevier Ltd. All rights reserved.

Can agroforest woodlots work as stepping stones for birds in the Atlantic forest region?

In fragmented landscapes, agroforest woodlots can potentially act as stepping stones facilitating movement between forest fragments. We assessed the influence of agroforest woodlots on bird distribution and diversity in the Atlantic forest region (SE Brazil), and also tested which categories of species can use different types of connection elements, and whether this use is influenced by the distance to large forest patches. We studied two fragmented landscapes, with and without stepping stones linking large fragments, and one forested landscape. Using a point count, a bird survey was undertaken in the fragmented landscapes in five different elements: large remnants (> 400 ha), agroforest woodlots (0.4-1.1 ha), small patches (0.5-7 ha), riparian corridor, and pasture areas (the main matrix). Generalist and open-area species were commonly observed in the agroforest system or other connection elements, whereas only a few forest species were present in these connections. For the latter species, the distance of woodlots to large patches was essential to determine their richness and abundance. Based on our results and data from literature, we suggest that there is an optimal relationship between the permeability of the matrix and the efficiency of stepping stones, which occurs at intermediate degrees of matrix resistance, and is species-dependent. Because the presence of agroforest system favors a higher richness of generalist species, they appeared to be more advantageous for conservation than the monoculture system; for this reason, they should be considered as a management alternative, particularly when the matrix permeability requirement is met.

Time-lag in biological responses to landscape changes in a highly dynamic Atlantic forest region

Time-lagged responses of biological variables to landscape modifications are widely recognized, but rarely considered in ecological studies. In order to test for the existence of time-lags in the response of trees, small mammals, birds and frogs to changes in fragment area and connectivity, we studied a fragmented and highly dynamic landscape in the Atlantic forest region. We also investigated the biological correlates associated with differential responses among taxonomic groups. Species richness and abundance for four taxonomic groups were measured in 21 secondary forest fragments during the same period (2000-2002), following a standardized protocol. Data analyses were based on power regressions and model selection procedures. The model inputs included present (2000) and past (1962, 1981) fragment areas and connectivity, as well as observed changes in these parameters. Although past landscape structure was particularly relevant for trees, all taxonomic groups (except small mammals) were affected by landscape dynamics, exhibiting a time-lagged response. Furthermore, fragment area was more important for species groups with lower dispersal capacity, while species with higher dispersal ability had stronger responses to connectivity measures. Although these secondary forest fragments still maintain a large fraction of their original biodiversity, the delay in biological response combined with high rates of deforestation and fast forest regeneration imply in a reduction in the average age of the forest. This also indicates that future species losses are likely, especially those that are more strictly-forest dwellers. Conservation actions should be implemented to reduce species extinction, to maintain old-growth forests and to favour the regeneration process. Our results demonstrate that landscape history can strongly affect the present distribution pattern of species in fragmented landscapes, and should be considered in conservation planning. (C) 2009 Elsevier Ltd. All rights reserved.

STRUCTURE OF THE SHRUB-ARBOREAL COMPONENT OF AN ATLANTIC FOREST FRAGMENT ON A HILLOCK IN THE CENTRAL LOWLAND OF RIO DE JANEIRO, BRAZIL

The present study describes and evaluates the horizontal and vertical structures of a lowland forest fragment on a hillock in the municipality of Silva Jardim, Rio de Janeiro State, Brazil (22 degrees 31`56 `` S and 42 degrees 20`46 `` W). Twenty plots (10x2m) totaling 0.5ha were laid out following the slope grade using DBH >= 5cm as the inclusion criterion. A total of 734 individuals were encountered, yielding a total density of 1468 ind./ha and a total basal area of 10783m(2). The richness values (129 species/41 families), Shannon-Wiener diversity (4.22) and equitability (0.87) indices indicated an accentuated floristic heterogeneity and low ecological dominance. Lauraceae, Myrtaceae, Fabaceae and Euphorbiaceae showed the greatest species richness, corroborating other studies that indicated these species as the most representative of Atlantic Forest areas in southeastern Brazil. The species with the greatest importance values (VI) were Aparisthmium cordatum, Guapira opposita, Lacistema pubescens, Xylopia sericea, Tapirira guianensis and Piptocarpha macropoda. The high diversity observed was influenced by earlier anthropogenic actions and by the current successional stage. The forest fragment studied demonstrated closer floristic similarity to areas inventoried in a close-by biological reserve than to fragments dispersed throughout the coastal plain. Similarities in soil type, degree of soil saturation and use-history of forest resources all support these relationships. The fragmented physiognomy of the central lowland in this region and the use-history of the landscape make these small remnant forest areas important in terms of establishing strategies for landscape restoration and species conservation.

Landscape and farm scale management to enhance biodiversity conservation in the cocoa producing region of southern Bahia, Brazil

In southern Bahia, Brazil, large land areas are used for the production of cocoa (Theobroma cacao), which is predominantly grown under the shade of native trees in an agroforestry system locally known as cabruca. As a dominant forest-like landscape element of the cocoa region, the cabrucas play an important role in the conservation of the region`s biodiversity. The purpose of this review is to provide the scientific basis for an action plan to reconcile cocoa production and biodiversity conservation in southern Bahia. The available research collectively highlights the diversity of responses of different species and biological groups to both the habitat quality of the cabrucas themselves and to the general characteristics of the landscape, such as the relative extent and spatial configuration of different vegetation types within the landscape mosaic. We identify factors that influence directly or indirectly the occurrence of native species in the cabrucas and the wider landscape of the cocoa region and develop recommendations for their conservation management. We show that the current scientific knowledge already provides a good basis for a biodiversity friendly management of the cocoa region of southern Bahia, although more work is needed to refine some management recommendations, especially on shade canopy composition and density, and verify their economic viability. The implementation of our recommendations should be accompanied by appropriate biological and socioeconomic monitoring and the findings should inform a broad program of adaptive management of the cabrucas and the wider cocoa landscape.

The spider genus Chrysometa (Araneae, Tetragnathidae) from the Pico da Neblina and Serra do Tapirapeco mountains (Amazonas, Brazil): new species, new records, diversity and distribution along two altitudinal gradients

Eight new species of the spider genus Chrysometa Simon, 1894 (Araneae, Tetragnathidae) are described and illustrated. Chrysometa nubigena n. sp., C. waikoxi n. sp., C. petrasierwaldae n. sp., C. santosi n. sp., C. yanomami n. sp., C. candianii n. sp., C. lomanhungae n. sp., and C. saci n. sp. Those species were collected in a study on the diversity of spider communities along altitudinal gradients in Brazilian Amazonia. C. saci was captured at the Serra do Tapirapeco (Barcelos), while all the other species are from the Pico da Neblina (Sao Gabriel da Cachoeira), the highest mountain in Brazil. We provide new records for C. boraceia, C. flava, C. guttata, C. minuta and C. opulenta, and we describe the male of C. minuta for the first time. We also present the first results on the diversity and altitudinal distribution of the species of Chrysometa at the Pico da Neblina and Serra do Tapirapeco. We sampled the first locality at six different elevations, and obtained 336 specimens distributed in 12 species. Richness and abundance, as well as relative importance peaked at the highest sites sampled (2,000 and 2,400 m). The three most abundant species showed a segregated distribution, being dominant or exclusively distributed in different altitudes. At the Serra do Tapirapeco, sampling at four different elevations up to 1200 m, we only obtained 40 individuals divided in four species, and there was no clear relation to altitude. Most of the new species were found at mid and high altitude sites, while species from lower altitude sites represented widespread species. The comparison with other neotropical spiders inventories highlights the high diversity recorded at Pico da Neblina, which could be assigned to the large environmental variation covered in this work and to the sampling of high-altitude environments. Inventories in the Andean region and other information in the literature also seem to support the association of Chrysometa with high altitude environments.

Conservation in tropical landscape mosaics: the case of the cacao landscape of southern Bahia, Brazil

A recent debate has contrasted two conservation strategies in agricultural landscapes; either ""land sparing`` farm development combining intensive production practices with forest set-asides, or ""wildlife-friendly`` farming with greater on-farm habitat value but lower yields. We argue that in established mosaic landscapes including old cacao production regions where natural forest has already been reduced to relatively small fragments, a combination of both strategies is needed to conserve biodiversity. After reviewing the evidence for the insufficiency of either strategy alone if applied to such landscapes, the paper focuses on the cacao production landscape of southern Bahia, Brazil, once the world`s second largest cacao producer. Here, small remaining areas of Atlantic Forest are embedded in a matrix dominated by traditional cacao agroforests, resulting in a landscape mosaic that has proven favorable to the conservation of the region`s high biodiversity. We show that current land use dynamics and public policies pose threats but also offer opportunities to conservation and describe a three- pronged landscape conservation strategy, consisting of (i) expansion of the protected areas system, (ii) promotion of productive yet biodiversity-friendly cacao farming practices, and (iii) assistance to land users to implement legally mandated on-farm reserves and voluntary private reserves. We discuss recent experiences concerning the implementation of this strategy, discuss likely future scenarios, and reflect on the applicability of the Bahian experience to biodiversity rich cacao production regions elsewhere in the tropics.