Disentangling the numbers behind agriculture-driven tropical deforestation.

Tropical deforestation continues at alarming rates with profound impacts on ecosystems, climate, and livelihoods, prompting renewed commitments to halt its continuation. Although it is well established that agriculture is a dominant driver of deforestation, rates and mechanisms remain disputed and often lack a clear evidence base. We synthesize the best available pantropical evidence to provide clarity on how agriculture drives deforestation. Although most (90 to 99%) deforestation across the tropics 2011 to 2015 was driven by agriculture, only 45 to 65% of deforested land became productive agriculture within a few years. Therefore, ending deforestation likely requires combining measures to create deforestation-free supply chains with landscape governance interventions. We highlight key remaining evidence gaps including deforestation trends, commodity-specific land-use dynamics, and data from tropical dry forests and forests across Africa.

A Commodity Supply Mix for More Regionalized Life Cycle Assessments.

Supply chain information is invaluable to further regionalize product life cycle assessments (LCAs), but detailed information linking production and consumption centers is not always available. We introduce the commodity supply mix (CSM) defined as the trade-volume-weighted average representing the combined geographic areas for the production of a commodity exported to a given market with the goal of (1) enhancing the relevance of inventory and impact regionalization and (2) allocating these impacts to specific markets. We apply the CSM to the Brazilian soybean supply chain mapped by Trase to obtain the mix of ecoregions and river basins linked to domestic consumption and exports to China, EU, France, and the rest of the world, before quantifying damage to biodiversity, and water scarcity footprints. The EU had the lowest potential biodiversity damage but the largest water scarcity footprint following respective sourcing patterns in 12 ecoregions and 18 river basins. These results differed from the average impact scores obtained from Brazilian soybean production information alone. The CSM can be derived at different scales (subnationally, internationally) using existing supply chain information and constitutes an additional step toward greater regionalization in LCAs, particularly for impacts with greater spatial variability such as biodiversity and water scarcity.

The timing and causality of ecological shifts on Caribbean reefs.

Caribbean reefs have experienced unprecedented changes in the past four decades. Of great concern is the perceived widespread shift from coral to macroalgal dominance and the question of whether it represents a new, stable equilibrium for coral-reef communities. The primary causes of the shift-grazing pressure (top-down), nutrient loading (bottom-up) or direct coral mortality (side-in)-still remain somewhat controversial in the coral-reef literature. We have attempted to tease out the relative importance of each of these causes. Four insights emerge from our analysis of an early regional dataset of information on the benthic composition of Caribbean reefs spanning the years 1977-2001. First, although three-quarters of reef sites have experienced coral declines concomitant with macroalgal increases, fewer than 10% of the more than 200 sites studied were dominated by macroalgae in 2001, by even the most conservative definition of dominance. Using relative dominance as the threshold, a total of 49 coral-to-macroalgae shifts were detected. This total represents ~35% of all sites that were dominated by coral at the start of their monitoring periods. Four shifts (8.2%) occurred because of coral loss with no change in macroalgal cover, 15 (30.6%) occurred because of macroalgal gain without coral loss, and 30 (61.2%) occurred owing to concomitant coral decline and macroalgal increase. Second, the timing of shifts at the regional scale is most consistent with the side-in model of reef degradation, which invokes coral mortality as a precursor to macroalgal takeover, because more shifts occurred after regional coral-mortality events than expected by chance. Third, instantaneous observations taken at the start and end of the time-series for individual sites showed these reefs existed along a continuum of coral and macroalgal cover. The continuous, broadly negative relationship between coral and macroalgal cover suggests that in some cases coral-to-macroalgae phase shifts may be reversed by removing sources of perturbation or restoring critical components such as the herbivorous sea urchin Diadema antillarum to the system. The five instances in which macroalgal dominance was reversed corroborate the conclusion that macroalgal dominance is not a stable, alternative community state as has been commonly assumed. Fourth, the fact that the loss in regional coral cover and concomitant changes to the benthic community are related to punctuated, discrete events with known causes (i.e. coral disease and bleaching), lends credence to the hypothesis that coral reefs of the Caribbean have been under assault from climate-change-related maladies since the 1970s.

Integrated terrestrial-freshwater planning doubles conservation of tropical aquatic species.

Conservation initiatives overwhelmingly focus on terrestrial biodiversity, and little is known about the freshwater cobenefits of terrestrial conservation actions. We sampled more than 1500 terrestrial and freshwater species in the Amazon and simulated conservation for species from both realms. Prioritizations based on terrestrial species yielded on average just 22% of the freshwater benefits achieved through freshwater-focused conservation. However, by using integrated cross-realm planning, freshwater benefits could be increased by up to 600% for a 1% reduction in terrestrial benefits. Where freshwater biodiversity data are unavailable but aquatic connectivity is accounted for, freshwater benefits could still be doubled for negligible losses of terrestrial coverage. Conservation actions are urgently needed to improve the status of freshwater species globally. Our results suggest that such gains can be achieved without compromising terrestrial conservation goals.

Author Correction: Strategic approaches to restoring ecosystems can triple conservation gains and halve costs.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Linking global drivers of agricultural trade to on-the-ground impacts on biodiversity.

Consumption of globally traded agricultural commodities like soy and palm oil is one of the primary causes of deforestation and biodiversity loss in some of the world's most species-rich ecosystems. However, the complexity of global supply chains has confounded efforts to reduce impacts. Companies and governments with sustainability commitments struggle to understand their own sourcing patterns, while the activities of more unscrupulous actors are conveniently masked by the opacity of global trade. We combine state-of-the-art material flow, economic trade, and biodiversity impact models to produce an innovative approach for understanding the impacts of trade on biodiversity loss and the roles of remote markets and actors. We do this for the production of soy in the Brazilian Cerrado, home to more than 5% of the world´s species. Distinct sourcing patterns of consumer countries and trading companies result in substantially different impacts on endemic species. Connections between individual buyers and specific hot spots explain the disproportionate impacts of some actors on endemic species and individual threatened species, such as the particular impact of European Union consumers on the recent habitat losses for the iconic giant anteater (Myrmecophaga tridactyla). In making these linkages explicit, our approach enables commodity buyers and investors to target their efforts much more closely to improve the sustainability of their supply chains in their sourcing regions while also transforming our ability to monitor the impact of such commitments over time.

Strategic approaches to restoring ecosystems can triple conservation gains and halve costs.

International commitments for ecosystem restoration add up to one-quarter of the world's arable land. Fulfilling them would ease global challenges such as climate change and biodiversity decline but could displace food production and impose financial costs on farmers. Here, we present a restoration prioritization approach capable of revealing these synergies and trade-offs, incorporating ecological and economic efficiencies of scale and modelling specific policy options. Using an actual large-scale restoration target of the Atlantic Forest hotspot, we show that our approach can deliver an eightfold increase in cost-effectiveness for biodiversity conservation compared with a baseline of non-systematic restoration. A compromise solution avoids 26% of the biome's current extinction debt of 2,864 plant and animal species (an increase of 257% compared with the baseline). Moreover, this solution sequesters 1 billion tonnes of CO2-equivalent (a 105% increase) while reducing costs by US$28 billion (a 57% decrease). Seizing similar opportunities elsewhere would offer substantial contributions to some of the greatest challenges for humankind.

Second rate or a second chance? Assessing biomass and biodiversity recovery in regenerating Amazonian forests.

Secondary forests (SFs) regenerating on previously deforested land account for large, expanding areas of tropical forest cover. Given that tropical forests rank among Earth's most important reservoirs of carbon and biodiversity, SFs play an increasingly pivotal role in the carbon cycle and as potential habitat for forest biota. Nevertheless, their capacity to regain the biotic attributes of undisturbed primary forests (UPFs) remains poorly understood. Here, we provide a comprehensive assessment of SF recovery, using extensive tropical biodiversity, biomass, and environmental datasets. These data, collected in 59 naturally regenerating SFs and 30 co-located UPFs in the eastern Amazon, cover >1,600 large- and small-stemmed plant, bird, and dung beetles species and a suite of forest structure, landscape context, and topoedaphic predictors. After up to 40 years of regeneration, the SFs we surveyed showed a high degree of biodiversity resilience, recovering, on average among taxa, 88% and 85% mean UPF species richness and composition, respectively. Across the first 20 years of succession, the period for which we have accurate SF age data, biomass recovered at 1.2% per year, equivalent to a carbon uptake rate of 2.25 Mg/ha per year, while, on average, species richness and composition recovered at 2.6% and 2.3% per year, respectively. For all taxonomic groups, biomass was strongly associated with SF species distributions. However, other variables describing habitat complexity-canopy cover and understory stem density-were equally important occurrence predictors for most taxa. Species responses to biomass revealed a successional transition at approximately 75 Mg/ha, marking the influx of high-conservation-value forest species. Overall, our results show that naturally regenerating SFs can accumulate substantial amounts of carbon and support many forest species. However, given that the surveyed SFs failed to return to a typical UPF state, SFs are not substitutes for UPFs.

The future of hyperdiverse tropical ecosystems.

The tropics contain the overwhelming majority of Earth's biodiversity: their terrestrial, freshwater and marine ecosystems hold more than three-quarters of all species, including almost all shallow-water corals and over 90% of terrestrial birds. However, tropical ecosystems are also subject to pervasive and interacting stressors, such as deforestation, overfishing and climate change, and they are set within a socio-economic context that includes growing pressure from an increasingly globalized world, larger and more affluent tropical populations, and weak governance and response capacities. Concerted local, national and international actions are urgently required to prevent a collapse of tropical biodiversity.

Disentangling the pathways of land use impacts on the functional structure of fish assemblages in Amazon streams.

Agricultural land use is a primary driver of environmental impacts on streams. However, the causal processes that shape these impacts operate through multiple pathways and at several spatial scales. This complexity undermines the development of more effective management approaches, and illustrates the need for more in-depth studies to assess the mechanisms that determine changes in stream biodiversity. Here we present results of the most comprehensive multi-scale assessment of the biological condition of streams in the Amazon to date, examining functional responses of fish assemblages to land use. We sampled fish assemblages from two large human-modified regions, and characterized stream conditions by physical habitat attributes and key landscape-change variables, including density of road crossings (i.e. riverscape fragmentation), deforestation, and agricultural intensification. Fish species were functionally characterized using ecomorphological traits describing feeding, locomotion, and habitat preferences, and these traits were used to derive indices that quantitatively describe the functional structure of the assemblages. Using structural equation modeling, we disentangled multiple drivers operating at different spatial scales, identifying causal pathways that significantly affect stream condition and the structure of the fish assemblages. Deforestation at catchment and riparian network scales altered the channel morphology and the stream bottom structure, changing the functional identity of assemblages. Local deforestation reduced the functional evenness of assemblages (i.e. increased dominance of specific trait combinations) mediated by expansion of aquatic vegetation cover. Riverscape fragmentation reduced functional richness, evenness and divergence, suggesting a trend toward functional homogenization and a reduced range of ecological niches within assemblages following the loss of regional connectivity. These results underscore the often-unrecognized importance of different land use changes, each of which can have marked effects on stream biodiversity. We draw on the relationships observed herein to suggest priorities for the improved management of stream systems in the multiple-use landscapes that predominate in human-modified tropical forests.

Is environmental legislation conserving tropical stream faunas? A large-scale assessment of local, riparian and catchment-scale influences on Amazonian fish.

Agricultural expansion and intensification are major threats to tropical biodiversity. In addition to the direct removal of native vegetation, agricultural expansion often elicits other human-induced disturbances, many of which are poorly addressed by existing environmental legislation and conservation programmes. This is particularly true for tropical freshwater systems, where there is considerable uncertainty about whether a legislative focus on protecting riparian vegetation is sufficient to conserve stream fauna.To assess the extent to which stream fish are being effectively conserved in agricultural landscapes, we examined the spatial distribution of assemblages in river basins to identify the relative importance of human impacts at instream, riparian and catchment scales, in shaping observed patterns. We used an extensive dataset on the ecological condition of 83 low-order streams distributed in three river basins in the eastern Brazilian Amazon.We collected and identified 24,420 individual fish from 134 species. Multiplicative diversity partitioning revealed high levels of compositional dissimilarity (DS) among stream sites (DS = 0.74 to 0.83) and river basins (DS = 0.82), due mainly to turnover (77.8% to 81.8%) rather than nestedness. The highly heterogeneous fish faunas in small Amazonian streams underscore the vital importance of enacting measures to protect forests on private lands outside of public protected areas.Instream habitat features explained more variability in fish assemblages (15%-19%) than riparian (2%-12%), catchment (4%-13%) or natural covariates (4%-11%). Although grouping species into functional guilds allowed us to explain up to 31% of their abundance (i.e. for nektonic herbivores), individual riparian - and catchment - scale predictor variables that are commonly a focus of environmental legislation explained very little of the observed variation (partial R2 values mostly <5%).Policy implications. Current rates of agricultural intensification and mechanization in tropical landscapes are unprecedented, yet the existing legislative frameworks focusing on protecting riparian vegetation seem insufficient to conserve stream environments and their fish assemblages. To safeguard the species-rich freshwater biota of small Amazonian streams, conservation actions must shift towards managing whole basins and drainage networks, as well as agricultural practices in already-cleared land.

Research priorities for managing the impacts and dependencies of business upon food, energy, water and the environment.

Delivering access to sufficient food, energy and water resources to ensure human wellbeing is a major concern for governments worldwide. However, it is crucial to account for the 'nexus' of interactions between these natural resources and the consequent implications for human wellbeing. The private sector has a critical role in driving positive change towards more sustainable nexus management and could reap considerable benefits from collaboration with researchers to devise solutions to some of the foremost sustainability challenges of today. Yet opportunities are missed because the private sector is rarely involved in the formulation of deliverable research priorities. We convened senior research scientists and influential business leaders to collaboratively identify the top forty questions that, if answered, would best help companies understand and manage their food-energy-water-environment nexus dependencies and impacts. Codification of the top order nexus themes highlighted research priorities around development of pragmatic yet credible tools that allow businesses to incorporate nexus interactions into their decision-making; demonstration of the business case for more sustainable nexus management; identification of the most effective levers for behaviour change; and understanding incentives or circumstances that allow individuals and businesses to take a leadership stance. Greater investment in the complex but productive relations between the private sector and research community will create deeper and more meaningful collaboration and cooperation.

Land use intensification in the humid tropics increased both alpha and beta diversity of soil bacteria.

Anthropogenic pressures on tropical forests are rapidly intensifying, but our understanding of their implications for biological diversity is still very limited, especially with regard to soil biota, and in particular soil bacterial communities. Here we evaluated bacterial community composition and diversity across a gradient of land use intensity in the eastern Amazon from undisturbed primary forest, through primary forests varyingly disturbed by fire, regenerating secondary forest, pasture, and mechanized agriculture. Soil bacteria were assessed by paired-end Illumina sequencing of 16S rRNA gene fragments (V4 region). The resulting sequences were clustered into operational taxonomic units (OTU) at a 97% similarity threshold. Land use intensification increased the observed bacterial diversity (both OTU richness and community heterogeneity across space) and this effect was strongly associated with changes in soil pH. Moreover, land use intensification and subsequent changes in soil fertility, especially pH, altered the bacterial community composition, with pastures and areas of mechanized agriculture displaying the most contrasting communities in relation to undisturbed primary forest. Together, these results indicate that tropical forest conversion impacts soil bacteria not through loss of diversity, as previously thought, but mainly by imposing marked shifts on bacterial community composition, with unknown yet potentially important implications for ecological functions and services performed by these communities.

Anthropogenic disturbance in tropical forests can double biodiversity loss from deforestation.

Concerted political attention has focused on reducing deforestation, and this remains the cornerstone of most biodiversity conservation strategies. However, maintaining forest cover may not reduce anthropogenic forest disturbances, which are rarely considered in conservation programmes. These disturbances occur both within forests, including selective logging and wildfires, and at the landscape level, through edge, area and isolation effects. Until now, the combined effect of anthropogenic disturbance on the conservation value of remnant primary forests has remained unknown, making it impossible to assess the relative importance of forest disturbance and forest loss. Here we address these knowledge gaps using a large data set of plants, birds and dung beetles (1,538, 460 and 156 species, respectively) sampled in 36 catchments in the Brazilian state of Pará. Catchments retaining more than 69­80% forest cover lost more conservation value from disturbance than from forest loss. For example, a 20% loss of primary forest, the maximum level of deforestation allowed on Amazonian properties under Brazil's Forest Code, resulted in a 39­54% loss of conservation value: 96­171% more than expected without considering disturbance effects. We extrapolated the disturbance-mediated loss of conservation value throughout Pará, which covers 25% of the Brazilian Amazon. Although disturbed forests retained considerable conservation value compared with deforested areas, the toll of disturbance outside Pará's strictly protected areas is equivalent to the loss of 92,000­139,000 km2 of primary forest. Even this lowest estimate is greater than the area deforested across the entire Brazilian Amazon between 2006 and 2015 (ref. 10). Species distribution models showed that both landscape and within-forest disturbances contributed to biodiversity loss, with the greatest negative effects on species of high conservation and functional value. These results demonstrate an urgent need for policy interventions that go beyond the maintenance of forest cover to safeguard the hyper-diversity of tropical forest ecosystems.

Idiosyncratic responses of Amazonian birds to primary forest disturbance.

As humans continue to alter tropical landscapes across the world, it is important to understand what environmental factors help determine the persistence of biodiversity in modified ecosystems. Studies on well-known taxonomic groups can offer critical insights as to the fate of biodiversity in these modified systems. Here we investigated species-specific responses of 44 forest-associated bird species with different behavioural traits to forest disturbance in 171 transects distributed across 31 landscapes in two regions of the eastern Brazilian Amazon. We investigated patterns of species occurrence in primary forests varyingly disturbed by selective-logging and fire and examined the relative importance of local, landscape and historical environmental variables in determining species occurrences. Within undisturbed and disturbed primary forest transects, we found that distance to forest edge and the biomass of large trees were the most important predictors driving the occurrence of individual species. However, we also found considerable variation in species responses to different environmental variables as well as inter-regional variation in the responses of the same species to the same environmental variables. We advocate the utility of using species-level analyses to complement community-wide responses in order to uncover highly variable and species-specific responses to environmental change that remain so poorly understood.

How pervasive is biotic homogenization in human-modified tropical forest landscapes?

Land-cover change and ecosystem degradation may lead to biotic homogenization, yet our understanding of this phenomenon over large spatial scales and different biotic groups remains weak. We used a multi-taxa dataset from 335 sites and 36 heterogeneous landscapes in the Brazilian Amazon to examine the potential for landscape-scale processes to modulate the cumulative effects of local disturbances. Biotic homogenization was high in production areas but much less in disturbed and regenerating forests, where high levels of among-site and among-landscape ß-diversity appeared to attenuate species loss at larger scales. We found consistently high levels of ß-diversity among landscapes for all land cover classes, providing support for landscape-scale divergence in species composition. Our findings support concerns that ß-diversity has been underestimated as a driver of biodiversity change and underscore the importance of maintaining a distributed network of reserves, including remaining areas of undisturbed primary forest, but also disturbed and regenerating forests, to conserve regional biota.

Developing Cost-Effective Field Assessments of Carbon Stocks in Human-Modified Tropical Forests.

Across the tropics, there is a growing financial investment in activities that aim to reduce emissions from deforestation and forest degradation, such as REDD+. However, most tropical countries lack on-the-ground capacity to conduct reliable and replicable assessments of forest carbon stocks, undermining their ability to secure long-term carbon finance for forest conservation programs. Clear guidance on how to reduce the monetary and time costs of field assessments of forest carbon can help tropical countries to overcome this capacity gap. Here we provide such guidance for cost-effective one-off field assessments of forest carbon stocks. We sampled a total of eight components from four different carbon pools (i.e. aboveground, dead wood, litter and soil) in 224 study plots distributed across two regions of eastern Amazon. For each component we estimated survey costs, contribution to total forest carbon stocks and sensitivity to disturbance. Sampling costs varied thirty-one-fold between the most expensive component, soil, and the least, leaf litter. Large live stems (≥10 cm DBH), which represented only 15% of the overall sampling costs, was by far the most important component to be assessed, as it stores the largest amount of carbon and is highly sensitive to disturbance. If large stems are not taxonomically identified, costs can be reduced by a further 51%, while incurring an error in aboveground carbon estimates of only 5% in primary forests, but 31% in secondary forests. For rapid assessments, necessary to help prioritize locations for carbon- conservation activities, sampling of stems ≥20cm DBH without taxonomic identification can predict with confidence (R2 = 0.85) whether an area is relatively carbon-rich or carbon-poor-an approach that is 74% cheaper than sampling and identifying all the stems ≥10cm DBH. We use these results to evaluate the reliability of forest carbon stock estimates provided by the IPCC and FAO when applied to human-modified forests, and to highlight areas where cost savings in carbon stock assessments could be most easily made.

Thresholds of species loss in Amazonian deforestation frontier landscapes.

In the Brazilian Amazon, private land accounts for the majority of remaining native vegetation. Understanding how land-use change affects the composition and distribution of biodiversity in farmlands is critical for improving conservation strategies in the face of rapid agricultural expansion. Working across an area exceeding 3 million ha in the southwestern state of Rondônia, we assessed how the extent and configuration of remnant forest in replicate 10,000-ha landscapes has affected the occurrence of a suite of Amazonian mammals and birds. In each of 31 landscapes, we used field sampling and semistructured interviews with landowners to determine the presence of 28 large and medium sized mammals and birds, as well as a further 7 understory birds. We then combined results of field surveys and interviews with a probabilistic model of deforestation. We found strong evidence for a threshold response of sampled biodiversity to landscape level forest cover; landscapes with <30-40% forest cover hosted markedly fewer species. Results from field surveys and interviews yielded similar thresholds. These results imply that in partially deforested landscapes many species are susceptible to extirpation following relatively small additional reductions in forest area. In the model of deforestation by 2030 the number of 10,000-ha landscapes under a conservative threshold of 43% forest cover almost doubled, such that only 22% of landscapes would likely to be able to sustain at least 75% of the 35 focal species we sampled. Brazilian law requires rural property owners in the Amazon to retain 80% forest cover, although this is rarely achieved. Prioritizing efforts to ensure that entire landscapes, rather than individual farms, retain at least 50% forest cover may help safeguard native biodiversity in private forest reserves in the Amazon.

Actor-specific contributions to the deforestation slowdown in the Brazilian Amazon.

Annual deforestation rates in the Brazilian Amazon fell by 77% between 2004 and 2011, yet have stabilized since 2009 at 5,000-7,000 km(2). We provide the first submunicipality assessment, to our knowledge, of actor-specific contributions to the deforestation slowdown by linking agricultural census and remote-sensing data on deforestation and forest degradation. Almost half (36,158 km(2)) of the deforestation between 2004 and 2011 occurred in areas dominated by larger properties (>500 ha), whereas only 12% (9,720 km(2)) occurred in areas dominated by smallholder properties (<100 ha). In addition, forests in areas dominated by smallholders tend to be less fragmented and less degraded. However, although annual deforestation rates fell during this period by 68-85% for all actors, the contribution of the largest landholders (>2,500 ha) to annual deforestation decreased over time (63% decrease between 2005 and 2011), whereas that of smallholders went up by a similar amount (69%) during the same period. In addition, the deforestation share attributable to remote areas increased by 88% between 2009 and 2011. These observations are consistent across the Brazilian Amazon, regardless of geographical differences in actor dominance or socioenvironmental context. Our findings suggest that deforestation policies to date, which have been particularly focused on command and control measures on larger properties in deforestation hotspots, may be increasingly limited in their effectiveness and fail to address all actors equally. Further reductions in deforestation are likely to be increasingly costly and require actor-tailored approaches, including better monitoring to detect small-scale deforestation and a shift toward more incentive-based conservation policies.

Road networks predict human influence on Amazonian bird communities.

Road building can lead to significant deleterious impacts on biodiversity, varying from direct road-kill mortality and direct habitat loss associated with road construction, to more subtle indirect impacts from edge effects and fragmentation. However, little work has been done to evaluate the specific effects of road networks and biodiversity loss beyond the more generalized effects of habitat loss. Here, we compared forest bird species richness and composition in the municipalities of Santarém and Belterra in Pará state, eastern Brazilian Amazon, with a road network metric called 'roadless volume (RV)' at the scale of small hydrological catchments (averaging 3721 ha). We found a significant positive relationship between RV and both forest bird richness and the average number of unique species (species represented by a single record) recorded at each site. Forest bird community composition was also significantly affected by RV. Moreover, there was no significant correlation between RV and forest cover, suggesting that road networks may impact biodiversity independently of changes in forest cover. However, variance partitioning analysis indicated that RV has partially independent and therefore additive effects, suggesting that RV and forest cover are best used in a complementary manner to investigate changes in biodiversity. Road impacts on avian species richness and composition independent of habitat loss may result from road-dependent habitat disturbance and fragmentation effects that are not captured by total percentage habitat cover, such as selective logging, fire, hunting, traffic disturbance, edge effects and road-induced fragmentation.