Linear infrastructure such as roads, highways, power lines and gas lines are omnipresent features of human activity and are rapidly expanding in the tropics. Tropical species are especially ...vulnerable to such infrastructure because they include many ecological specialists that avoid even narrow (<30-m wide) clearings and forest edges, as well as other species that are susceptible to road kill, predation or hunting by humans near roads. In addition, roads have a major role in opening up forested tropical regions to destructive colonization and exploitation. Here, we synthesize existing research on the impacts of roads and other linear clearings on tropical rainforests, and assert that such impacts are often qualitatively and quantitatively different in tropical forests than in other ecosystems. We also highlight practical measures to reduce the negative impacts of roads and other linear infrastructure on tropical species.
ABSTRACT
We synthesize findings from one of the world's largest and longest‐running experimental investigations, the Biological Dynamics of Forest Fragments Project (BDFFP). Spanning an area of ∼1000 ...km2 in central Amazonia, the BDFFP was initially designed to evaluate the effects of fragment area on rainforest biodiversity and ecological processes. However, over its 38‐year history to date the project has far transcended its original mission, and now focuses more broadly on landscape dynamics, forest regeneration, regional‐ and global‐change phenomena, and their potential interactions and implications for Amazonian forest conservation. The project has yielded a wealth of insights into the ecological and environmental changes in fragmented forests. For instance, many rainforest species are naturally rare and hence are either missing entirely from many fragments or so sparsely represented as to have little chance of long‐term survival. Additionally, edge effects are a prominent driver of fragment dynamics, strongly affecting forest microclimate, tree mortality, carbon storage and a diversity of fauna.
Even within our controlled study area, the landscape has been highly dynamic: for example, the matrix of vegetation surrounding fragments has changed markedly over time, succeeding from large cattle pastures or forest clearcuts to secondary regrowth forest. This, in turn, has influenced the dynamics of plant and animal communities and their trajectories of change over time. In general, fauna and flora have responded differently to fragmentation: the most locally extinction‐prone animal species are those that have both large area requirements and low tolerance of the modified habitats surrounding fragments, whereas the most vulnerable plants are those that respond poorly to edge effects or chronic forest disturbances, and that rely on vulnerable animals for seed dispersal or pollination.
Relative to intact forests, most fragments are hyperdynamic, with unstable or fluctuating populations of species in response to a variety of external vicissitudes. Rare weather events such as droughts, windstorms and floods have had strong impacts on fragments and left lasting legacies of change. Both forest fragments and the intact forests in our study area appear to be influenced by larger‐scale environmental drivers operating at regional or global scales. These drivers are apparently increasing forest productivity and have led to concerted, widespread increases in forest dynamics and plant growth, shifts in tree‐community composition, and increases in liana (woody vine) abundance. Such large‐scale drivers are likely to interact synergistically with habitat fragmentation, exacerbating its effects for some species and ecological phenomena. Hence, the impacts of fragmentation on Amazonian biodiversity and ecosystem processes appear to be a consequence not only of local site features but also of broader changes occurring at landscape, regional and even global scales.
ABSTRACT
Increased frequency and severity of drought, as a result of climate change, is expected to drive critical changes in plant–insect interactions that may elevate rates of tree mortality. The ...mechanisms that link water stress in plants to insect performance are not well understood. Here, we build on previous reviews and develop a framework that incorporates the severity and longevity of drought and captures the plant physiological adjustments that follow moderate and severe drought. Using this framework, we investigate in greater depth how insect performance responds to increasing drought severity for: (i) different feeding guilds; (ii) flush feeders and senescence feeders; (iii) specialist and generalist insect herbivores; and (iv) temperate versus tropical forest communities. We outline how intermittent and moderate drought can result in increases of carbon‐based and nitrogen‐based chemical defences, whereas long and severe drought events can result in decreases in plant secondary defence compounds. We predict that different herbivore feeding guilds will show different but predictable responses to drought events, with most feeding guilds being negatively affected by water stress, with the exception of wood borers and bark beetles during severe drought and sap‐sucking insects and leaf miners during moderate and intermittent drought. Time of feeding and host specificity are important considerations. Some insects, regardless of feeding guild, prefer to feed on younger tissues from leaf flush, whereas others are adapted to feed on senescing tissues of severely stressed trees. We argue that moderate water stress could benefit specialist insect herbivores, while generalists might prefer severe drought conditions. Current evidence suggests that insect outbreaks are shorter and more spatially restricted in tropical than in temperate forests. We suggest that future research on the impact of drought on insect communities should include (i) assessing how drought‐induced changes in various plant traits, such as secondary compound concentrations and leaf water potential, affect herbivores; (ii) food web implications for other insects and those that feed on them; and (iii) interactions between the effects on insects of increasing drought and other forms of environmental change including rising temperatures and CO2 levels. There is a need for larger, temperate and tropical forest‐scale drought experiments to look at herbivorous insect responses and their role in tree death.
Edge effects are major drivers of change in many fragmented landscapes, but are often highly variable in space and time. Here we assess variability in edge effects altering Amazon forest dynamics, ...plant community composition, invading species, and carbon storage, in the world's largest and longest-running experimental study of habitat fragmentation. Despite detailed knowledge of local landscape conditions, spatial variability in edge effects was only partially foreseeable: relatively predictable effects were caused by the differing proximity of plots to forest edge and varying matrix vegetation, but windstorms generated much random variability. Temporal variability in edge phenomena was also only partially predictable: forest dynamics varied somewhat with fragment age, but also fluctuated markedly over time, evidently because of sporadic droughts and windstorms. Given the acute sensitivity of habitat fragments to local landscape and weather dynamics, we predict that fragments within the same landscape will tend to converge in species composition, whereas those in different landscapes will diverge in composition. This 'landscape-divergence hypothesis', if generally valid, will have key implications for biodiversity-conservation strategies and for understanding the dynamics of fragmented ecosystems.
We synthesize findings to date from the world’s largest and longest-running experimental study of habitat fragmentation, located in central Amazonia. Over the past 32
years, Amazonian forest ...fragments ranging from 1 to 100
ha have experienced a wide array of ecological changes. Edge effects have been a dominant driver of fragment dynamics, strongly affecting forest microclimate, tree mortality, carbon storage, fauna, and other aspects of fragment ecology. However, edge-effect intensity varies markedly in space and time, and is influenced by factors such as edge age, the number of nearby edges, and the adjoining matrix of modified vegetation surrounding fragments. In our study area, the matrix has changed markedly over the course of the study (evolving from large cattle pastures to mosaics of abandoned pasture and regrowth forest) and this in turn has strongly influenced fragment dynamics and faunal persistence. Rare weather events, especially windstorms and droughts, have further altered fragment ecology. In general, populations and communities of species in fragments are hyperdynamic relative to nearby intact forest. Some edge and fragment-isolation effects have declined with a partial recovery of secondary forests around fragments, but other changes, such as altered patterns of tree recruitment, are ongoing. Fragments are highly sensitive to external vicissitudes, and even small changes in local land-management practices may drive fragmented ecosystems in markedly different directions. The effects of fragmentation are likely to interact synergistically with other anthropogenic threats such as logging, hunting, and especially fire, creating an even greater peril for the Amazonian biota.
Forest fragmentation is considered a greater threat to vertebrates than to tree communities because individual trees are typically long-lived and require only small areas for survival. Here we show ...that forest fragmentation provokes surprisingly rapid and profound alterations in Amazonian tree-community composition. Results were derived from a 22-year study of exceptionally diverse tree communities in 40 1-ha plots in fragmented and intact forests, which were sampled repeatedly before and after fragment isolation. Within these plots, trajectories of change in abundance were assessed for 267 genera and 1,162 tree species. Abrupt shifts in floristic composition were driven by sharply accelerated tree mortality and recruitment within approximately 100 m of fragment margins, causing rapid species turnover and population declines or local extinctions of many large-seeded, slow-growing, and old-growth taxa; a striking increase in a smaller set of disturbance-adapted and abiotically dispersed species; and significant shifts in tree size distributions. Even among old-growth trees, species composition in fragments is being restructured substantially, with subcanopy species that rely on animal seed-dispersers and have obligate outbreeding being the most strongly disadvantaged. These diverse changes in tree communities are likely to have wide-ranging impacts on forest architecture, canopy-gap dynamics, plant-animal interactions, and forest carbon storage.
Understanding how tropical rainforest trees may respond to the precipitation extremes predicted in future climate change scenarios is paramount for their conservation and management. Tree species ...clearly differ in drought susceptibility, suggesting that variable water transport strategies exist. Using a multi-disciplinary approach, we examined the hydraulic variability in trees in a lowland tropical rainforest in north-eastern Australia. We studied eight tree species representing broad plant functional groups (one palm and seven eudicot mature-phase, and early-successional trees). We characterised the species' hydraulic system through maximum rates of volumetric sap flow and velocities using the heat ratio method, and measured rates of tree growth and several stem, vessel, and leaf traits. Sap flow measures exhibited limited variability across species, although early-successional species and palms had high mean sap velocities relative to most mature-phase species. Stem, vessel, and leaf traits were poor predictors of sap flow measures. However, these traits exhibited different associations in multivariate analysis, revealing gradients in some traits across species and alternative hydraulic strategies in others. Trait differences across and within tree functional groups reflect variation in water transport and drought resistance strategies. These varying strategies will help in our understanding of changing species distributions under predicted drought scenarios.
We synthesized key findings from the Biological Dynamics of Forest Fragments Project, the world's largest and longest-running experimental study of habitat fragmentation. Although initially designed ...to assess the influence of fragment area on Amazonian biotas, the project has yielded insights that go far beyond the original scope of the study. Results suggest that edge effects play a key role in fragment dynamics, that the matrix has a major influence on fragment connectivity and functioning, and that many Amazonian species avoid even small (<100-m-wide) clearings. The effects of fragmentation are highly eclectic, altering species richness and abundances, species invasions, forest dynamics, the trophic structure of communities, and a variety of ecological and ecosystem processes. Moreover, forest fragmentation appears to interact synergistically with ecological changes such as hunting, fires, and logging, collectively posing an even greater threat to the rainforest biota.
The impacts of potential linear barriers such as roads, highways, and power lines on rainforest fauna are poorly understood. In the central Brazilian Amazon, we compared the frequency of local ...movements (≤300 m long) of understory birds within intact forest and across a 30- to 40-m-wide road over a 2-year period. Rainforest had regenerated along some road verges, to the extent that a nearly complete canopy was formed in some areas, so we also assessed whether this facilitated bird movement. Movement data were determined from 1212 recaptures of 3681 netted birds at six study sites. The road significantly inhibited total bird movement across roads at five of the six sites. Bird foraging guilds varied in their responses to the road and different ages of regrowth. Movements of frugivorous and edge and gap species were not inhibited at any site, whereas most forest-dependent insectivores (mixed-species flocks, terrestrial species, and army-ant followers) had markedly inhibited road-crossing movements, except at sites with extensive regrowth. Solitary understory species were especially vulnerable, rarely crossing even roads overgrown by tall regrowth. For sensitive species, road-crossing movements were inhibited because individuals tended to avoid both edge-affected habitat near the road and the road clearing itself. Our results suggest that even narrow roads with low traffic volumes can reduce local movements of many insectivorous birds in Amazonia.
Assessing the persistent impacts of fragmentation on aboveground structure of tropical forests is essential to understanding the consequences of land use change for carbon storage and other ecosystem ...functions. We investigated the influence of edge distance and fragment size on canopy structure, aboveground woody biomass (AGB), and AGB turnover in the Biological Dynamics of Forest Fragments Project (BDFFP) in central Amazon, Brazil, after 22+ yr of fragment isolation, by combining canopy variables collected with portable canopy profiling lidar and airborne laser scanning surveys with long-term forest inventories. Forest height decreased by 30% at edges of large fragments (>10 ha) and interiors of small fragments (<3 ha). In larger fragments, canopy height was reduced up to 40 m from edges. Leaf area density profiles differed near edges: the density of understory vegetation was higher and midstory vegetation lower, consistent with canopy reorganization via increased regeneration of pioneers following post-fragmentation mortality of large trees. However, canopy openness and leaf area index remained similar to control plots throughout fragments, while canopy spatial heterogeneity was generally lower at edges. AGB stocks and fluxes were positively related to canopy height and negatively related to spatial heterogeneity. Other forest structure variables typically used to assess the ecological impacts of fragmentation (basal area, density of individuals, and density of pioneer trees) were also related to lidar-derived canopy surface variables. Canopy reorganization through the replacement of edge-sensitive species by disturbance-tolerant ones may have mitigated the biomass loss effects due to fragmentation observed in the earlier years of BDFFP. Lidar technology offered novel insights and observational scales for analysis of theecological impacts of fragmentation on forest structure and function, specifically aboveground biomass storage.
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