Linking changes in landscape structure to insect herbivory in forest edges and interiors of Atlantic Forest remnants.

Human activities have triggered profound changes in natural landscapes, resulting in species loss and disruption of pivotal ecological interactions such as insect herbivory. This antagonistic interaction is affected by complex pathways (e.g., abundance of herbivores and predators, plant chemical defenses, and resource availability), but the knowledge regarding how forest loss and fragmentation affect insect herbivory in human-modified tropical landscapes still remains poorly understood. In this context, we assessed multi-pathways by which changes in landscape structure likely influence insect herbivory in 20 Atlantic forest fragments in Brazil. Using path analysis, we estimated the direct effects of forest cover and forest edge density, and the indirect effect via canopy openness, number of understory plants and phenolic compounds, on leaf damage in understory plants located in the edge and interior of forest fragments. In particular, plants located in forest edges experienced greater leaf damage than interior ones. We observed that landscape edge density exerted a positive and direct effect on leaf damage in plants sampled at the edge of forest fragments. Our findings also indicated that forest loss and increase of edge density led to an increase in the canopy opening in the forest interior, which causes a reduction in the number of understory plants and, consequently, an increase in leaf damage. In addition, we detected that phenolic compounds negatively influence leaf damage in forest interior plants. Given the increasing forest loss in tropical regions, in which forest fragments become stranded in highly deforested, edge-dominated and degraded landscapes, our study highlights the pervasive enhancement in insect herbivory in remaining forest fragments-especially along forest edges and canopy gaps in the forest interior. As a result, increased herbivory is likely to affect forest regeneration and accelerate the ecological meltdown processes in these highly deforested and disturbed anthropogenic landscapes.

Land-use homogenization reduces the occurrence and diversity of frugivorous birds in a tropical biodiversity hotspot.

Understanding how human-modified landscapes maintain biodiversity and provide ecosystem services is crucial for establishing conservation practices. Given that responses to land-use are species-specific, it is crucial to understand how land-use changes may shape patterns of species diversity and persistence in human-modified landscapes. Here, we used a comprehensive data set on bird distribution from the Brazilian Atlantic Forest to understand how species richness and individual occurrences of frugivorous bird species responded to land-use spatial predictors and, subsequently, assess how ecological traits and phylogeny modulated these responses. Using Bayesian hierarchical modeling, we reveal that the richness of frugivorous birds was positively associated with the amount of native forest and negatively with both agriculture and pasture amount at the landscape scale. Conversely, the effect of these predictors on species occurrence and ecological traits was highly variable and presented a weak phylogenetic signal. Furthermore, land-use homogenization (i.e., the conversion of forest to pasture or agriculture) led to pervasive consequences for forest-dependent bird species, whereas several generalist species thrived in deforested areas, replacing those sensitive to habitat disturbances.

Preserving 40% forest cover is a valuable and well-supported conservation guideline: reply to Banks-Leite et al.

Banks-Leite et al. (2021) claim that our suggestion of preserving ≥ 40% forest cover lacks evidence and can be problematic. We find these claims unfounded, and discuss why conservation planning urgently requires valuable, well-supported and feasible general guidelines like the 40% criterion. Using region-specific thresholds worldwide is unfeasible and potentially harmful.

Determinants of population persistence and abundance of terrestrial and arboreal vertebrates stranded in tropical forest land-bridge islands.

Megadams are among the key modern drivers of habitat and biodiversity loss in emerging economies. The Balbina Hydroelectric Dam of Central Brazilian Amazonia inundated 312,900 ha of primary forests and created approximately 3500 variable-sized islands that still harbor vertebrate populations after nearly 3 decades after isolation. We estimated the species richness, abundance, biomass, composition, and group size of medium- to large-bodied forest vertebrates in response to patch, landscape, and habitat-quality metrics across 37 islands and 3 continuous forest sites throughout the Balbina archipelago. We conducted 1168 km of diurnal censuses and had 12,420 camera-trapping days along 81 transects with 207 camera stations. We determined the number of individuals (or groups) detected per 10 km walked and the number of independent photographs per 10 camera-trapping days, respectively, for each species. We recorded 34 species, and patch area was the most significant predictor of vertebrate population relative abundance and aggregate biomass. The maximum group size of several group-living species was consistently larger on large islands and in continuous patches than on small islands. Most vertebrate populations were extirpated after inundation. Remaining populations are unlikely to survive further ecological disruptions. If all vertebrate species were once widely distributed before inundation, we estimated that approximately 75% of all individual vertebrates were lost from all 3546 islands and 7.4% of the animals in all persisting insular populations are highly likely to be extirpated. Our results demonstrate that population abundance estimates should be factored into predictions of community disassembly on small islands to robustly predict biodiversity outcomes. Given the rapidly escalating hydropower infrastructure projects in developing counties, we suggest that faunal abundance and biomass estimates be considered in environmental impact assessments and large strictly protected reserves be established to minimize detrimental effects of dams on biodiversity. Conserving large tracts of continuous forests represents the most critical conservation measure to ensure that animal populations can persist at natural densities in Amazonian forests.

Determinantes de la Persistencia Poblacional y la Abundancia de Vertebrados Terrestres y Arbóreos Varados en Islas con Puentes Terrestres en los Bosques Tropicales Resumen Las megapresas se encuentran entre los causantes modernos de la pérdida de hábitat y biodiversidad en las economías emergentes. La Presa Hidroeléctrica Balbina de la Amazonia central en Brasil inundó 312, 900 ha de bosques primarios y creó ∼3500 islas de tamaños variables que todavía albergan poblaciones de vertebrados después de casi tres décadas de aislamiento. Estimamos la riqueza de especies, abundancia, biomasa, composición y tamaño grupal de los vertebrados de talla mediana a grande del bosque como respuesta a la a las medidas de calidad del fragmento, paisaje y hábitat en 37 islas y tres sitios de bosque continuo en todo el archipiélago de Balbina. Realizamos censos diurnos a lo largo de 1,168 km y tuvimos 12,420 días de trampeo con cámaras a lo largo de 81 transectos con 207 estaciones de cámara. Determinamos el número de individuos (o grupos) detectados en cada diez kilómetros recorridos y el número de fotografías independientes para cada diez días de trampeo con cámara, respectivamente, para cada especie. Registramos 34 especies y el área del fragmento fue el pronosticador más importante de la población de vertebrados en relación con la abundancia y la biomasa agregada. El tamaño grupal máximo de varias de las especies gregarias fue sistemáticamente mayor en las islas grandes y en los fragmentos continuos que en las islas pequeñas. La mayoría de las poblaciones de vertebrados fueron extirpadas después de la inundación. Las poblaciones remanentes tienen pocas probabilidades de sobrevivir más perturbaciones ecológicas. Si todas las especies de vertebrados tuvieron alguna vez una distribución amplia previa a la inundación, estimamos que se perdió ∼75% de todos los vertebrados individuales en todas las 3, 546 islas y el 7.4% de los animales en las poblaciones insulares remanentes tienen una alta probabilidad de desaparecer. Nuestros resultados demuestran que las estimaciones de abundancia poblacional deberían incluirse en las predicciones del desmontaje comunitario en islas pequeñas para predecir con solidez los resultados de biodiversidad. Ya que cada vez existen más proyectos de infraestructura hidroeléctrica en los países en desarrollo, sugerimos que las estimaciones de abundancia de fauna y de biomasa sean consideradas en las evaluaciones de impacto ambiental y que se establezcan grandes reservas con protección estricta para minimizar los efectos dañinos que tienen las presas sobre la biodiversidad. La conservación de grandes tramos continuos de bosque representa la medida de conservación más crítica para asegurar que las poblaciones animales puedan persistir con densidades naturales en los bosques de la Amazonia.

Habitat loss and canopy openness mediate leaf trait plasticity of an endangered palm in the Brazilian Atlantic Forest.

Forest cover and light availability comprise key factors for plant establishment in tropical forests. In the Brazilian Atlantic Forest (AF), Euterpe edulis (Areacaceae) is an endangered and keystone food resource contributing to forest functionality. We investigated the influence of forest loss and light availability on leaf traits and acclimatization of young individuals of E. edulis in AF fragments. We aimed to understand (i) how canopy openness and transmitted light are affected by forest cover at the landscape scale and the individual palm level; and (ii) how local and landscape features, combined and separately, affect key leaf traits widely known to be related to plant growth. The study was carried out in 15 forest fragments, ranging from 16 to 97% of surrounding forest cover. In each fragment, we sampled 10-20 individuals of E. edulis and analyzed nine leaf traits related to morphological, biochemical and chemical aspects. We also took hemispherical photographs to estimate canopy openness on the top of each E. edulis and also within fragment plots. We found that young plants predominantly occurred in more shaded environments. Additionally, E. edulis succeeded to acclimate in six of the nine traits analyzed, with most traits being affected by local and landscape features. It is likely that the lack of variation in traits related to protection against herbivory are limiting the species establishment in highly deforested landscapes. Our results provide novel evidence that both landscape and local contexts affect the leaf traits of E. edulis young plants leading to biochemical, chemical and morphological adjustments.

Designing optimal human-modified landscapes for forest biodiversity conservation.

Agriculture and development transform forest ecosystems to human-modified landscapes. Decades of research in ecology have generated myriad concepts for the appropriate management of these landscapes. Yet, these concepts are often contradictory and apply at different spatial scales, making the design of biodiversity-friendly landscapes challenging. Here, we combine concepts with empirical support to design optimal landscape scenarios for forest-dwelling species. The supported concepts indicate that appropriately sized landscapes should contain ≥ 40% forest cover, although higher percentages are likely needed in the tropics. Forest cover should be configured with c. 10% in a very large forest patch, and the remaining 30% in many evenly dispersed smaller patches and semi-natural treed elements (e.g. vegetation corridors). Importantly, the patches should be embedded in a high-quality matrix. The proposed landscape scenarios represent an optimal compromise between delivery of goods and services to humans and preserving most forest wildlife, and can therefore guide forest preservation and restoration strategies.

Small mammal responses to Amazonian forest islands are modulated by their forest dependence.

Hydroelectric dams have induced widespread loss, fragmentation and degradation of terrestrial habitats in lowland tropical forests. Yet their ecological impacts have been widely neglected, particularly in developing countries, which are currently earmarked for exponential hydropower development. Here we assess small mammal assemblage responses to Amazonian forest habitat insularization induced by the 28-year-old Balbina Hydroelectric Dam. We sampled small mammals on 25 forest islands (0.83-1466 ha) and four continuous forest sites in the mainland to assess the overall community structure and species-specific responses to forest insularization. We classified all species according to their degree of forest-dependency using a multi-scale approach, considering landscape, patch and local habitat characteristics. Based on 65,520 trap-nights, we recorded 884 individuals of at least 22 small mammal species. Species richness was best predicted by island area and isolation, with small islands (< 15 ha) harbouring an impoverished nested subset of species (mean ± SD: 2.6 ± 1.3 species), whereas large islands (> 200 ha; 10.8 ± 1.3 species) and continuous forest sites (∞ ha; 12.5 ± 2.5 species) exhibited similarly high species richness. Forest-dependent species showed higher local extinction rates and were often either absent or persisted at low abundances on small islands, where non-forest-dependent species became hyper-abundant. Species capacity to use non-forest habitat matrices appears to dictate small mammal success in small isolated islands. We suggest that ecosystem functioning may be highly disrupted on small islands, which account for 62.7% of all 3546 islands in the Balbina Reservoir.

Lessons from a Community-Based Program to Monitor Forest Vertebrates in the Brazilian Amazon.

A large number of sustainable use reserves recently have been titled in the Brazilian Amazonia. These reserves require public participation in the design and implementation of management and monitoring programs. Species-monitoring programs that engage local stakeholders may be useful for assessing wildlife status over the long term. We collaborated on the development of a participatory program to monitor forest vertebrates in the Piagaçu-Purus Sustainable Development Reserve and to build capacity among the local people. We examined relations between the distance to the nearest human community and sighting rates of each species, and evaluated the program overall. Eighteen wildlife monitors received training in line transect and sign surveys and then conducted surveys along a total of ten transects. Sighting rates of most species in the Piagaçu-Purus Sustainable Development Reserve were higher than those reported in other Amazonian forests. Distance to the human community was not associated with the overall vertebrate sighting rate. Use of the trained monitors was successful in terms of data acquisition and engagement. The involvement of local people promoted discussions about regulation of hunting in the reserve. Implementation of community-based programs to monitor forest wildlife in Amazonian sustainable use reserves may empower local communities and assess the status of wildlife through time.

Predicting primate local extinctions within "real-world" forest fragments: a pan-neotropical analysis.

Understanding the main drivers of species extinction in human-modified landscapes has gained paramount importance in proposing sound conservation strategies. Primates play a crucial role in maintaining the integrity of forest ecosystem functions and represent the best studied order of tropical terrestrial vertebrates, yet primate species diverge widely in their responses to forest habitat disturbance and fragmentation. Here, we present a robust quantitative review on the synergistic effects of habitat fragmentation on Neotropical forest primates to pinpoint the drivers of species extinction across a wide range of forest patches from Mexico to Argentina. Presence-absence data on 19 primate functional groups were compiled from 705 forest patches and 55 adjacent continuous forest sites, which were nested within 61 landscapes investigated by 96 studies. Forest patches were defined in terms of their size, surrounding matrix and level of hunting pressure on primates, and each functional group was classified according to seven life-history traits. Generalized linear mixed models showed that patch size, forest cover, level of hunting pressure, home range size and trophic status were the main predictors of species persistence within forest isolates for all functional groups pooled together. However, patterns of local extinction varied greatly across taxa, with Alouatta and Callicebus moloch showing the highest occupancy rates even within tiny forest patches, whereas Brachyteles and Leontopithecus occupied fewer than 50% of sites, even in relatively large forest tracts. Our results uncover the main predictors of platyrrhine primate species extinction, highlighting the importance of considering the history of anthropogenic disturbances, the structure of landscapes, and species life-history attributes in predicting primate persistence in Neotropical forest patches. We suggest that large-scale conservation planning of fragmented forest landscapes should prioritize and set-aside large, well-connected and strictly protected forest reserves to maximize species persistence across the entire spectrum of primate life-history.

A New Species of Nealiolus Mason, 1974 (Hymenoptera: Braconidae) from Traditional Cocoa Agroforestry and Brazilian Atlantic Rainforest.

The genus Nealiolus Mason, 1974 (Hymenoptera: Braconidae, Brachistinae) is recorded for the first time in the Atlantic forest remnants and traditional cocoa agroforests, Theobroma cacao (Malvaceae) in the northeastern Brazilian region. A new species Nealiolus seb Silva, Shimbori & Fernandes sp. n. is described and named in honor of the "Sociedade Entomológica do Brasil - SEB", on the occasion of its fiftieth anniversary (golden jubilee). Digital images, map and an identification key for all Brazilian species of Nealiolus are provided.

Terrestrial food web complexity in Amazonian forests decays with habitat loss.

The conversion of natural ecosystems into human-modified landscapes (HMLs) is the main driver of biodiversity loss in terrestrial ecosystems.1,2,3 Even when species persist within habitat remnants, populations may become so small that ecological interactions are functionally lost, disrupting local interaction networks.4,5 To uncover the consequences of land use changes toward ecosystem functioning, we need to understand how changes in species richness and abundance in HMLs6,7,8 rearrange ecological networks. We used data from forest vertebrate surveys and combined modeling and network analysis to investigate how the structure of predator-prey networks was affected by habitat insularization induced by a hydroelectric reservoir in the Brazilian Amazonia.9 We found that network complexity, measured by interaction diversity, decayed non-linearly with decreasingly smaller forest area. Although on large forest islands (>100 ha) prey species were linked to 3-4 potential predators, they were linked to one or had no remaining predator on small islands. Using extinction simulations, we show that the variation in network structure cannot be explained by abundance-related extinction risk or prey availability. Our findings show that habitat loss may result in an abrupt disruption of terrestrial predator-prey networks, generating low-complexity ecosystems that may not retain functionality. Release from predation on some small islands may produce cascading effects over plants that accelerate forest degradation, whereas predator spillover on others may result in overexploited prey populations. Our analyses highlight that in addition to maintaining diversity, protecting large continuous forests is required for the persistence of interaction networks and related ecosystem functions.

Emergent properties of species-habitat networks in an insular forest landscape.

Deforestation and fragmentation are pervasive drivers of biodiversity loss, but how they scale up to entire landscapes remains poorly understood. Here, we apply species-habitat networks based on species co-occurrences to test the effects of insular fragmentation on multiple taxa-medium-large mammals, small nonvolant mammals, lizards, understory birds, frogs, dung beetles, orchid bees, and trees-across 22 forest islands and three continuous forest sites within a river-damming quasi-experimental landscape in Central Amazonia. Widespread, nonrandom local species extinctions were translated into highly nested networks of low connectance and modularity. Networks' robustness considering the sequential removal of large-to-small sites was generally low; between 5% (dung beetles) and 50% (orchid bees) of species persisted when retaining only <10 ha of islands. In turn, larger sites and body size were the main attributes structuring the networks. Our results raise the prospects that insular forest fragmentation results in simplified species-habitat networks, with distinct taxa persistence to habitat loss.

Gene Flow and Genetic Structure Reveal Reduced Diversity between Generations of a Tropical Tree, Manilkara multifida Penn., in Atlantic Forest Fragments.

The Atlantic Forest remnants in southern Bahia, Brazil, contain large tree species that have suffered disturbances in recent decades. Anthropogenic activities have led to a decrease in the population of many tree species and a loss of alleles that can maintain the evolutionary fitness of their populations. This study assessed patterns of genetic diversity, spatial genetic structure, and genetic structure among Manilkara multifida Penn. populations, comparing the genetic parameters of adult and juvenile trees. In particular, we collected leaves from adults and juveniles of M. multifida in two protected areas, the Veracel Station (EVC) and the Una Biological Reserve (UBR), located in threatened Atlantic Forest fragments. We observed a substantial decay in genetic variability between generations in both areas i.e., adults' HO values were higher (EVC = 0.720, UBR = 0.736) than juveniles' (EVC = 0.463 and UBR = 0.560). Both juveniles and adults showed genetic structure between the two areas (θ = 0.017 for adults and θ = 0.109 for juveniles). Additionally, forest fragments indicated an unexpectedly short gene flow. Our results, therefore, highlight the pervasive effects of historical deforestation and other human disturbances on the genetic diversity of M. multifida populations within a key conservation region of the Atlantic Forest biodiversity hotspot.

Woody lianas increase in dominance and maintain compositional integrity across an Amazonian dam-induced fragmented landscape.

Tropical forest fragmentation creates insular biological communities that undergo species loss and changes in community composition over time, due to area- and edge-effects. Woody lianas thrive in degraded and secondary forests, due to their competitive advantage over trees in these habitats. Lianas compete both directly and indirectly with trees, increasing tree mortality and turnover. Despite our growing understanding of liana-tree dynamics, we lack detailed knowledge of the assemblage-level responses of lianas themselves to fragmentation, particularly in evergreen tropical forests. We examine the responses of both sapling and mature liana communities to landscape-scale forest insularization induced by a mega hydroelectric dam in the Brazilian Amazon. Detailed field inventories were conducted on islands created during reservoir filling, and in nearby mainland continuous forest. We assess the relative importance of variables associated with habitat fragmentation such as area, isolation, surrounding forest cover, fire and wind disturbance, on liana community attributes including abundance, basal area, diversity, and composition. We also explore patterns of liana dominance relative to tree saplings and adults ≥10 cm diameter at breast height. We find that 1) liana community composition remains remarkably similar across mainland continuous forest and islands, regardless of extreme area- and edge- effects and the loss of vertebrate dispersers in the latter; and 2) lianas are increasing in dominance relative to trees in the sapling layer in the most degraded islands, with both the amount of forest cover surrounding islands and fire disturbance history predicting liana dominance. Our data suggest that liana communities persist intact in isolated forests, regardless of extreme area- and edge-effects; while in contrast, tree communities simultaneously show evidence of increased turnover and supressed recruitment. These processes may lead to lianas becoming a dominant component of this dam-induced fragmented landscape in the future, due to their competitive advantage over trees in degraded forest habitats. Additional loss of tree biomass and diversity brought about through competition with lianas, and the concurrent loss of carbon storage, should be accounted for in impact assessments of future dam development.

Functional decay in tree community within tropical fragmented landscapes: Effects of landscape-scale forest cover.

As tropical rainforests are cleared, forest remnants are increasingly isolated within agricultural landscapes. Understanding how forest loss impacts on species diversity can, therefore, contribute to identifying the minimum amount of habitat required for biodiversity maintenance in human-modified landscapes. Here, we evaluate how the amount of forest cover, at the landscape scale, affects patterns of species richness, abundance, key functional traits and common taxonomic families of adult trees in twenty Brazilian Atlantic rainforest landscapes. We found that as forest cover decreases, both tree community richness and abundance decline, without exhibiting a threshold. At the family-level, species richness and abundance of the Myrtaceae and Sapotaceae were also negatively impacted by the percent forest remaining at the landscape scale. For functional traits, we found a reduction in shade-tolerant, animal-dispersed and small-seeded species following a decrease in the amount of forest retained in landscapes. These results suggest that the amount of forest in a landscape is driving non-random losses in phylogenetic and functional tree diversity in Brazil's remaining Atlantic rainforests. Our study highlights potential restraints on the conservation value of Atlantic rainforest remnants in deforested landscapes in the future.

Widespread Forest Vertebrate Extinctions Induced by a Mega Hydroelectric Dam in Lowland Amazonia.

Mega hydropower projects in tropical forests pose a major emergent threat to terrestrial and freshwater biodiversity worldwide. Despite the unprecedented number of existing, under-construction and planned hydroelectric dams in lowland tropical forests, long-term effects on biodiversity have yet to be evaluated. We examine how medium and large-bodied assemblages of terrestrial and arboreal vertebrates (including 35 mammal, bird and tortoise species) responded to the drastic 26-year post-isolation history of archipelagic alteration in landscape structure and habitat quality in a major hydroelectric reservoir of Central Amazonia. The Balbina Hydroelectric Dam inundated 3,129 km2 of primary forests, simultaneously isolating 3,546 land-bridge islands. We conducted intensive biodiversity surveys at 37 of those islands and three adjacent continuous forests using a combination of four survey techniques, and detected strong forest habitat area effects in explaining patterns of vertebrate extinction. Beyond clear area effects, edge-mediated surface fire disturbance was the most important additional driver of species loss, particularly in islands smaller than 10 ha. Based on species-area models, we predict that only 0.7% of all islands now harbor a species-rich vertebrate assemblage consisting of ≥80% of all species. We highlight the colossal erosion in vertebrate diversity driven by a man-made dam and show that the biodiversity impacts of mega dams in lowland tropical forest regions have been severely overlooked. The geopolitical strategy to deploy many more large hydropower infrastructure projects in regions like lowland Amazonia should be urgently reassessed, and we strongly advise that long-term biodiversity impacts should be explicitly included in pre-approval environmental impact assessments.