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1. Habitat loss is the primary cause of local extinctions. Yet, there is considerable uncertainty regarding how fast species respond to habitat loss, and how time-delayed responses vary in space. 2. We focused on the Argentine Dry Chaco (c. 32 million ha), a global deforestation hotspot, and tested for time-delayed response of bird and mammal communities to landscape transformation. We quantified the magnitude of extinction debt by modelling contemporary species richness as a function of either contemporary or past (2000 and 1985) landscape patterns. We then used these models to map communities' extinction debt. 3. We found strong evidence for an extinction debt: landscape structure from 2000 explained contemporary species richness of birds and mammals better than contemporary and 1985 landscapes. This suggests time-delayed responses between 10 and 25 years. Extinction debt was especially strong for forest specialists. 4. Projecting our models across the Chaco highlighted areas where future local extinctions due to unpaid extinction debt are likely. Areas recently converted to agriculture had highest extinction debt, regardless of the post-conversion land use. Few local extinctions were predicted in areas with remaining larger forest patches. 5. Synthesis and applications. The evidence for an unpaid extinction debt in the Argentine Dry Chaco provides a substantial window of opportunity for averting local biodiversity losses. However, this window may close rapidly if conservation activities such as habitat restoration are not implemented swiftly. Our extinction debt maps highlight areas where such conservation activities should be implemented.

1. The fruit-tracking hypothesis predicts spatiotemporal links between changes in the abundance of fruit-eating birds and the abundance of their fleshy-fruit resources. 2. While the spatial scale of plant—frugivore interactions has been explored to understand mismatches between observed and expected fruit—frugivore patterns, methodological issues such as the consequences of measuring fruit and frugivore abundance rather than fruit availability and fruit consumption have not been evaluated. 3. Here, we explored whether predicted fruit—frugivore spatiotemporal links can be captured with higher accuracy by proximate measurements of interaction strength. We used a 6-ha grided plot in an Andean subtropical forest to study the link between (i) fruit and fruit-eating bird abundances; (ii) fruit availability and frequency of fruit consumption; and (iii) covariation between frugivore abundance and frequency of frugivory. We evaluated these links for the entire frugivore assemblage and for the four most important species using data gathered bimonthly along a 2-year period. 4. Fleshy-fruit availability and abundance varied sharply temporally and were patchily distributed in mosaics that differed in fruit quantity. Fruit availability and abundance also varied along spatial gradients extended over the whole study plot. We found a strong response of the entire frugivorous bird assemblage to fruit availability over time, and a weakly significant relationship over space at the local scale. The main frugivore species widely differed in their responses to changes in fruit abundance in such a way that response at the assemblage level cannot be seen as the sum of individual responses of each species. Our results suggest that fruit tracking in frugivorous—insectivorous birds may be largely explained by species-specific responses to changes in the availability of fruits and alternative resources. 5. In agreement with our prediction, more accurate measurements of interaction strength described fruit—frugivore relationships better than traditional measurements. Moreover, we show that covariation between frugivore abundance, frequency of fruit consumption and fruit availability must be included in the fruit-tracking hypothesis framework to demonstrate (or reject) spatiotemporal fruit tracking. We propose that estimation of nutrient and energy availability in fruits could be a new frontier to understanding the forces driving foraging decisions that lead to fruit tracking.

The multi‐scale spatial match between bird and food abundances is a main driver of the structure of fruit‐eating bird assemblages. We explored how the activity of fruit‐eating birds was influenced by the abundance of fruits at the local and landscape scales in Andean mountain forests during the breeding season, when most birds forage close to their nest. We measured: (1) the spatial scale of variation in the abundance of fruits, (2) the spatial scale of variation in the activity of fruit‐eating birds, and (3) the spatial match between both variables. The sampling design consisted of eleven 1.2‐ha sites, each subdivided into 30 cells of 20 × 20 m, where we sampled fruits and fruit‐eating birds. We found that fruit consumption, and to a lesser extent bird abundance, were associated with local spatial variation in abundance of selected fruit species. However, fruit‐eating birds did not modify their spatial distribution in the landscape following changes in availability of these fruits. Our study shows that fruit‐eating birds detect local spatial variation in fruit availability in their home breeding ranges, and exploit patches with large clusters of selected fruits. However, it may be unprofitable for breeding birds to stray too far from their nests to exploit fruit‐rich patches, accounting for the absence of fruit tracking at larger spatial scales.

The White-fronted Woodpecker (Melanerpes cactorum) drills holes in branches and trunks to feed on sap flows, providing an energy-rich food resource for other birds. Here we describe ecological and behavioral traits of the White-fronted Woodpecker related to its sap-feeding habits in the semiarid Chaco of Argentina and explore the structure of the avian assemblage in relation to the sap resource. Sap consumption by the White-fronted Woodpecker and other sap-feeding species was strongly seasonal and positively associated with periods of resource scarcity. The White-fronted Woodpecker actively defended the sap wells from smaller birds. Specialist and facultative nectarivores that assimilate sucrose at a high rate represented an important proportion of sap-feeding birds. In this system of woodpecker, sap, and other sap-feeding species, each species' consumption depends on its physiological and behavioral characteristics as well as on the availability of other food in the surrounding environment.

Understanding the trade-offs between biodiversity conservation and agricultural production has become a fundamental question in sustainability science. Substantial research has focused on how species’ populations respond to agricultural intensification, with the goal to understand whether conservation policies that spatially separate agriculture and conservation or, alternatively, integrate the two are more beneficial. Spatial heterogeneity in both species abundance and agricultural productivity have been largely left out of this discussion, although these patterns are ubiquitous from local to global scales due to varying land capacity. Here, we address the question of how to align agricultural production and biodiversity conservation in heterogeneous landscapes. Using model simulations of species abundance and agricultural yields, we show that trade-offs between agricultural production and species’ abundance can be reduced by minimizing the cost (in terms of species abundance) of agricultural production. We find that when species’ abundance and agricultural yields vary across landscapes, the optimal strategy to minimize trade-offs is rarely pure land sparing or land sharing. Instead, landscapes that combine elements of both strategies are optimal. Additionally, we show how the reference population of a species is defined has important influences on optimization results. Our findings suggest that in the real world, understanding the impact of heterogeneous land capacity on biodiversity and agricultural production is crucial to designing multi-use landscapes that jointly maximize conservation and agricultural benefits.

Land-use change is a root cause of the extinction crisis, but links between habitat change and biodiversity loss are not fully understood. While there is evidence that habitat loss is an important extinction driver, the relevance of habitat fragmentation remains debated. Moreover, while time delays of biodiversity responses to habitat transformation are well-documented, time-delayed effects have been ignored in the habitat loss versus fragmentation debate. Here, using a hierarchical Bayesian multi-species occupancy framework, we systematically tested for time-delayed responses of bird and mammal communities to habitat loss and to habitat fragmentation. We focused on the Argentine Chaco, where deforestation has been widespread recently. We used an extensive field dataset on birds and mammals, along with a time series of annual woodland maps from 1985 to 2016 covering recent and historical habitat transformations. Contemporary habitat amount explained bird and mammal occupancy better than past habitat amount. However, occupancy was affected more by the past rather than recent fragmentation, indicating a time-delayed response to fragmentation. Considering past landscape patterns is therefore crucial for understanding current biodiversity patterns. Not accounting for land-use history ignores the possibility of extinction debt and can thus obscure impacts of fragmentation, potentially explaining contrasting findings of habitat loss versus fragmentation studies.

World‐wide, tropical savannas and dry forests are under increasing pressure from land use. The environmental impacts of agricultural expansion into these ecosystems have received much attention, yet subtler changes in natural vegetation remain severely understudied. We explored how bird communities vary along gradients of woody vegetation in the South American Dry Chaco by combining high‐resolution, satellite‐based tree, shrub and total woody cover with field data on the frequency of 82 bird species surveyed in 167 plots. We identified change points along woody cover gradients where the relative frequency of individual bird species dropped most strongly. Based on this, we identified forest indicator species and assessed evidence for community‐level thresholds. Most forest birds (71%) had clear change points in their frequencies along vegetation gradients, starting as high as 38% total woody cover. Many (41%) forest species declined drastically at woody cover levels of less than 11%. This general pattern was similar for tree and shrub cover. Only 7% of our study area had woody cover levels where we detected no response in forest bird communities. In contrast, 68% of the area had woody cover levels with incremental declines in forest bird species, and 25% of the study area had woody cover levels below the forest bird community threshold. We identified 11 indicator species strongly related to woody cover, with highest frequencies in the eastern and western Dry Chaco. Spatial distributions of these species corresponded well with areas above and below woody vegetation thresholds. Synthesis and applications. We found evidence for critical thresholds for forest birds along woody cover gradients in dry forests and implemented tools to map where these thresholds have been crossed. For the Chaco, we highlight the importance of maintaining woody cover levels above about 40%, such as in certain silvopastoral systems that can be much more wildlife‐friendly than other forms of agriculture. We identify remaining areas of potentially intact forest bird communities. More broadly, this study highlights the value of combining species‐level (indicator species' distributions) and ecosystem‐level (satellite‐based, continuous woody cover maps) surrogates for understanding biodiversity patterns and threats. We found evidence for critical thresholds for forest birds along woody cover gradients in dry forests and implemented tools to map where these thresholds have been crossed. For the Chaco, we highlight the importance of maintaining woody cover levels above about 40%, such as in certain silvopastoral systems that can be much more wildlife‐friendly than other forms of agriculture. We identify remaining areas of potentially intact forest bird communities. More broadly, this study highlights the value of combining species‐level (indicator species' distributions) and ecosystem‐level (satellite‐based, continuous woody cover maps) surrogates for understanding biodiversity patterns and threats.

Fil: Jobbagy Gampel, Esteban Gabrie. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - San Luis. Instituto de Matemática Aplicada de San Luis ; Argentina

Land-use change is a root cause of the extinction crisis, but links between habitat change and biodiversity loss are not fully understood. While there is evidence that habitat loss is an important extinction driver, the relevance of habitat fragmentation remains debated. Moreover, while time delays of biodiversity responses to habitat transformation are well-documented, timedelayed effects have been ignored in the habitat loss versus fragmentation debate. Here, using a hierarchical Bayesian multi-species occupancy framework, we systematically tested for time-delayed responses of bird and mammal communities to habitat loss and to habitat fragmentation. We focused on the Argentine Chaco, where deforestation has been widespread recently. We used an extensive field dataset on birds and mammals, along with a time series of annual woodland maps from 1985 to 2016 covering recent and historical habitat transformations. Contemporary habitat amount explained bird andmammal occupancy better than past habitat amount. However, occupancy was affected more by the past rather than recent fragmentation, indicating a time-delayed response to fragmentation. Considering past landscape patterns is therefore crucial for understanding current biodiversity patterns. Not accounting for land-use history ignores the possibility of extinction debt and can thus obscure impacts of fragmentation, potentially explaining contrasting findings of habitat loss versus fragmentation studies.

The biodiversity impacts of agricultural deforestation vary widely across regions. Previous efforts to explain this variation have focused exclusively on the landscape features and management regimes of agricultural systems, neglecting the potentially critical role of ecological filtering in shaping deforestation tolerance of extant species assemblages at large geographical scales via selection for functional traits. Here we provide a large-scale test of this role using a global database of species abundance ratios between matched agricultural and native forest sites that comprises 71 avian assemblages reported in 44 primary studies, and a companion database of 10 functional traits for all 2,647 species involved. Using meta-analytic, phylogenetic and multivariate methods, we show that beyond agricultural features, filtering by the extent of natural environmental variability and the severity of historical anthropogenic deforestation shapes the varying deforestation impacts across species assemblages. For assemblages under greater environmental variability—proxied by drier and more seasonal climates under a greater disturbance regime—and longer deforestation histories, filtering has attenuated the negative impacts of current deforestation by selecting for functional traits linked to stronger deforestation tolerance. Our study provides a previously largely missing piece of knowledge in understanding and managing the biodiversity consequences of deforestation by agricultural deforestation. The authors collate literature on the responses of bird assemblages to forest loss and show that locations with a more variable natural environment and a longer history of agricultural land use have bird assemblages that are more tolerant to forest loss.