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  1. Abstract Background

    Understanding how to connect habitat remnants to facilitate the movement of species is a critical task in an increasingly fragmented world impacted by human activities. The identification of dispersal routes and corridors through connectivity analysis requires measures of landscape resistance but there has been no consensus on how to calculate resistance from habitat characteristics, potentially leading to very different connectivity outcomes.

    Methods

    We propose a new model, called the Time-Explicit Habitat Selection (TEHS) model, that can be directly used for connectivity analysis. The TEHS model decomposes the movement process in a principled approach into a time and a selection component, providing complementary information regarding space use by separately assessing the drivers of time to traverse the landscape and the drivers of habitat selection. These models are illustrated using GPS-tracking data from giant anteaters (Myrmecophaga tridactyla) in the Pantanal wetlands of Brazil.

    Results

    The time model revealed that the fastest movements tended to occur between 8 p.m. and 5 a.m., suggesting a crepuscular/nocturnal behavior. Giant anteaters moved faster over wetlands while moving much slower over forests and savannas, in comparison to grasslands. We also found that wetlands were consistently avoided whereas forest and savannas tended to be selected. Importantly, this model revealed that selection for forest increased with temperature, suggesting that forests may act as important thermal shelters when temperatures are high. Finally, using the spatial absorbing Markov chain framework, we show that the TEHS model results can be used to simulate movement and connectivity within a fragmented landscape, revealing that giant anteaters will often not use the shortest-distance path to the destination patch due to avoidance of certain habitats.

    Conclusions

    The proposed approach can be used to characterize how landscape features are perceived by individuals through the decomposition of movement patterns into a time and a habitat selection component. Additionally, this framework can help bridge the gap between movement-based models and connectivity analysis, enabling the generation of time-explicit connectivity results.

     
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  2. Abstract

    Habitat loss is often considered the greatest near‐term threat to biodiversity, while the impact of habitat fragmentation remains intensely debated. A key issue of this debate centers on the problem of scale–landscape or patch–at which to assess the consequences of fragmentation. Yet patterns are often confounded across scales, and experimental designs that could solve this scaling problem remain scarce. We conducted two field experiments in 30 experimental landscapes in which we manipulated habitat loss, fragmentation, and patch size for a community of four insect herbivores that specialize on the cactusOpuntia. In the first experiment, we destroyed 2088Opuntiapatches in either aggregated or random patterns and compared the relative effects of landscape‐scale loss and fragmentation to those of local patch size on species occurrence. This experiment focused on manipulating the relative separation of remaining patches, where we hypothesized that aggregated loss would disrupt dispersal more than random loss, leading to lower occurrence. In the second experiment, we destroyed 759Opuntiapatches to generate landscapes that varied in patch number and size for a given amount of habitat loss and assessed species occurrence. This experiment focused on manipulating the subdivision of remaining habitat, where we hypothesized that an increase in the number of patches for a given amount of loss would lead to negative effects on occurrence. For both, we expected that occurrence would increase with patch size. We find strong evidence for landscape‐scale effects of habitat fragmentation, with aggregated loss and a larger number of patches for a given amount of habitat loss leading to a lower frequency of patches occupied in landscapes. In both experiments, occurrence increased with patch size, yet interactions of patch size and landscape‐scale loss and fragmentation drove species occurrence in patches. Importantly, the direction of effects were consistent across scales and effects of patch size were sufficient to predict the effects of habitat loss and fragmentation across entire landscapes. Our experimental results suggest that changes at both the patch and landscape scales can impact populations, but that a long‐standing pattern—the patch‐size effect—captures much of the key variation shaping patterns of species occurrence.

     
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  3. Abstract Fragmentation and scale

    Although habitat loss has well‐known impacts on biodiversity, the effects of habitat fragmentation remain intensely debated. It is often argued that the effects of habitat fragmentation, or the breaking apart of habitat for a given habitat amount, can be understood only at the scale of entire landscapes composed of multiple habitat patches. Yet, fragmentation also impacts the size, isolation and habitat edge for individual patches within landscapes. Addressing the problem of scale on fragmentation effects is crucial for resolving how fragmentation impacts biodiversity.

    Scaling framework

    We build upon scaling concepts in ecology to describe a framework that emphasizes three “dimensions” of scale in habitat fragmentation research: the scales of phenomena (or mechanisms), sampling and analysis. Using this framework, we identify ongoing challenges and provide guidance for advancing the science of fragmentation.

    Implications

    We show that patch‐ and landscape‐scale patterns arising from habitat fragmentation for a given amount of habitat are fundamentally related, leading to interdependencies among expected patterns arising from different scales of phenomena. Aggregation of information when increasing the grain of sampling (e.g., from patch to landscape) creates challenges owing to biases created from the modifiable areal unit problem. Consequently, we recommend that sampling strategies use the finest grain that captures potential underlying mechanisms (e.g., plot or patch). Study designs that can capture phenomena operating at multiple spatial extents offer the most promise for understanding the effects of fragmentation and its underlying mechanisms. By embracing the interrelationships among scales, we expect more rapid advances in our understanding of habitat fragmentation.

     
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  4. Connectivity has long played a central role in ecological and evolutionary theory and is increasingly emphasized for conserving biodiversity. Nonetheless, connectivity assessments often focus on individual species even though understanding and preserving connectivity for entire communities is urgently needed. Here we derive and test a framework that harnesses the well-known allometric scaling of animal movement to predict community-level connectivity across protected area networks. We used a field translocation experiment involving 39 species of southern African birds to quantify movement capacity, scaled this relationship to realized dispersal distances determined from ring-and-recovery banding data, and used allometric scaling equations to quantify community-level connectivity based on multilayer network theory. The translocation experiment explained observed dispersal distances from ring-recovery data and emphasized allometric scaling of dispersal based on morphology. Our community-level networks predicted that larger-bodied species had a relatively high potential for connectivity, while small-bodied species had lower connectivity. These community networks explained substantial variation in observed bird diversity across protected areas. Our results highlight that harnessing allometric scaling can be an effective way of determining large-scale community connectivity. We argue that this trait-based framework founded on allometric scaling provides a means to predict connectivity for entire communities, which can foster empirical tests of community theory and contribute to biodiversity conservation strategies aimed at mitigating the effects of environmental change.

     
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  5. Abstract

    Understanding animal movement often relies upon telemetry and biologging devices. These data are frequently used to estimate latent behavioural states to help understand why animals move across the landscape. While there are a variety of methods that make behavioural inferences from biotelemetry data, some features of these methods (e.g. analysis of a single data stream, use of parametric distributions) may limit their generality to reliably discriminate among behavioural states.

    To address some of the limitations of existing behavioural state estimation models, we introduce a nonparametric Bayesian framework called the mixed‐membership method for movement (M4), which is available within the open‐sourcebayesmoveR package. This framework can analyse multiple data streams (e.g. step length, turning angle, acceleration) without relying on parametric distributions, which may capture complex behaviours more successfully than current methods. We tested our Bayesian framework using simulated trajectories and compared model performance against two segmentation methods (behavioural change point analysis (BCPA) and segclust2d), one machine learning method [expectation‐maximization binary clustering (EMbC)] and one type of state‐space model [hidden Markov model (HMM)]. We also illustrated this Bayesian framework using movements of juvenile snail kitesRostrhamus sociabilisin Florida, USA.

    The Bayesian framework estimated breakpoints more accurately than the other segmentation methods for tracks of different lengths. Likewise, the Bayesian framework provided more accurate estimates of behaviour than the other state estimation methods when simulations were generated from less frequently considered distributions (e.g. truncated normal, beta, uniform). Three behavioural states were estimated from snail kite movements, which were labelled as ‘encamped’, ‘area‐restricted search’ and ‘transit’. Changes in these behaviours over time were associated with known dispersal events from the nest site, as well as movements to and from possible breeding locations.

    Our nonparametric Bayesian framework estimated behavioural states with comparable or superior accuracy compared to the other methods when step lengths and turning angles of simulations were generated from less frequently considered distributions. Since the most appropriate parametric distributions may not be obvious a priori, methods (such as M4) that are agnostic to the underlying distributions can provide powerful alternatives to address questions in movement ecology.

     
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  6. Abstract

    Maintaining the ability of organisms to move between suitable patches of habitat despite ongoing habitat loss is essential to conserving biodiversity. Quantifying connectivity has therefore become a central focus of conservation planning. A large number of metrics have been developed to estimate potential connectivity based on habitat configuration, matrix structure and information on organismal movement, and it is often assumed that metrics explain actual connectivity. Yet, validation of metrics is rare, particularly across entire landscapes undergoing habitat loss—a crucial problem that connectivity conservation aims to mitigate.

    We leveraged a landscape‐scale habitat loss and fragmentation experiment to assess the performance of commonly used patch‐ and landscape‐scale connectivity metrics against observed movement data, test whether incorporating information about the matrix improves connectivity metrics and examine the performance of metrics across a gradient of habitat loss. We tested whether 38 connectivity metrics predict movement at the patch (i.e. patch immigration rates) and landscape (i.e., total movements) scale for a pest insect, the cactus bugChelinidea vittiger, across 15 replicate landscapes.

    Metrics varied widely in their ability to explain actual connectivity. At the patch scale, dPCflux, which describes the contribution of a patch to movement across the landscape independent of patch size, best explained immigration rates. At the landscape scale, total movements were best explained by a mesoscale metric that captures that distance between clusters of patches (i.e. modules). Incorporating the matrix did not necessarily improve the ability of metrics to predict actual connectivity. Across the habitat loss gradient, dPCfluxwas sensitive to habitat amount.

    Synthesis and applications. Our study provides a much‐needed evaluation of network connectivity metrics at the patch and landscape scales, emphasizing that accurate quantification of connectivity requires the incorporation, not only of habitat amount but also habitat configuration and information on dispersal capability of species. We suggest that variation in habitat may often be more critical for interpreting network connectivity than the matrix, and advise that connectivity metrics may be sensitive to habitat loss and should therefore be applied with caution to highly fragmented landscapes. Finally, we recommend that applications integrate mesoscale configuration of habitat into connectivity strategies.

     
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  7. Abstract

    Selective logging is the primary cause of tropical forest degradation and is rapidly expanding worldwide. While the impacts of logging on species diversity and distributions are well understood, little is known about the effects of logging on animal behaviours central to individual fitness and population persistence.

    The song rate of breeding songbirds is a behavioural trait that is often positively associated with male density and used by conspecific females as an indicator of territory and male quality. Thus, contrasting logging‐induced adjustments in song rates of individual birds with population shifts may illuminate potential mechanisms underlying population distributions.

    We present a novel application of bioacoustic monitoring, integrating counts of individuals, songs and duets from single automated recording units (ARUs) withN‐mixture models, to estimate shifts in population parameters (occupancy, abundance) and singing behaviours (per‐capita song rates, per‐pair duet rates) of 32 Bornean songbird species with logging. We tested hypotheses on the relationships between adjustments in behavioural and population parameters with logging, and further tested the extent to which species traits predicted behavioural and population shifts.

    Adjustments to singing behaviour in 59 and 53% of species (57% of duetting species) were concordant with differences in occupancy and abundance respectively, such that species showing reduced populations with logging also produced fewer songs per‐capita, and vice versa. Species known to prefer undisturbed habitats and large‐bodied species showed the most negative effects of logging on singing behaviour and population distributions. Species known to exploit degraded habitats exhibited the opposite pattern. Subdued singing in logged forests by species of conservation concern suggests limited competition between territorial males in small populations and may also signal low‐quality territories.

    Synthesis and applications. We demonstrate that bioacoustic monitoring can be used to not only estimate important population parameters of occupancy and abundance across a diverse tropical songbird community, but also enables quantification of behaviours considered relevant to individual fitness, yet unobtainable with conventional methods (e.g. point counts). Bioacoustics provides a viable approach to reliable automated large‐scale monitoring of hyperdiverse tropical forest systems under logging operations and other land‐use pressures.

     
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