Identifying potential provenances for climate-change adaptation using spatially variable coefficient models.

BACKGROUND: Selection of climate-change adapted ecotypes of commercially valuable species to date relies on DNA-assisted screening followed by growth trials. For trees, such trials can take decades, hence any approach that supports focussing on a likely set of candidates may save time and money. We use a non-stationary statistical analysis with spatially varying coefficients to identify ecotypes that indicate first regions of similarly adapted varieties of Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco) in North America. For over 70,000 plot-level presence-absences, spatial differences in the survival response to climatic conditions are identified. RESULTS: The spatially-variable coefficient model fits the data substantially better than a stationary, i.e. constant-effect analysis (as measured by AIC to account for differences in model complexity). Also, clustering the model terms identifies several potential ecotypes that could not be derived from clustering climatic conditions itself. Comparing these six identified ecotypes to known genetically diverging regions shows some congruence, as well as some mismatches. However, comparing ecotypes among each other, we find clear differences in their climate niches. CONCLUSION: While our approach is data-demanding and computationally expensive, with the increasing availability of data on species distributions this may be a useful first screening step during the search for climate-change adapted varieties. With our unsupervised learning approach being explorative, finely resolved genotypic data would be helpful to improve its quantitative validation.

Predicting plant-pollinator interactions: concepts, methods, and challenges.

Plant-pollinator interactions are ecologically and economically important, and, as a result, their prediction is a crucial theoretical and applied goal for ecologists. Although various analytical methods are available, we still have a limited ability to predict plant-pollinator interactions. The predictive ability of different plant-pollinator interaction models depends on the specific definitions used to conceptualize and quantify species attributes (e.g., morphological traits), sampling effects (e.g., detection probabilities), and data resolution and availability. Progress in the study of plant-pollinator interactions requires conceptual and methodological advances concerning the mechanisms and species attributes governing interactions as well as improved modeling approaches to predict interactions. Current methods to predict plant-pollinator interactions present ample opportunities for improvement and spark new horizons for basic and applied research.

Exotic tree species have consistently lower herbivore load in a cross-Atlantic tree biodiversity experiment.

It is commonly expected that exotic plants experience reduced herbivory, but experimental evidence for such enemy release is still controversial. One reason for conflicting results might be that community context has rarely been accounted for, although the surrounding plant diversity may moderate enemy release. Here, we tested the effects of focal tree origin and surrounding tree diversity on herbivore abundance and leaf damage in a cross-Atlantic tree-diversity experiment in Canada and Germany. We evaluated six European tree species paired with six North American congeners in both their native and exotic range, expecting lower herbivory for the exotic tree species in each pair at each site. Such reciprocal experiments have long been called for, but have not been realized thus far. In addition to a thorough evaluation of overall enemy release effects, we tested whether enemy release effects changed with the surrounding tree diversity. Herbivore abundance was indeed consistently lower on exotics across all six tree genera (12 comparisons). This effect of exotic status was independent of the continent, phylogenetic relatedness, and surrounding tree diversity. In contrast, leaf damage associated with generalist leaf chewers was consistently higher on North American tree species. Interestingly, several species of European weevils were the most abundant leaf chewers on both continents and the dominant herbivores at the Canadian site. Thus, most observed leaf damage is likely to reflect the effect of generalist herbivores that feed heavily on plant species with which they have not evolved. At the German site, sap suckers were the dominant herbivores and showed a pattern consistent with enemy release. Taken together, the consistently lower herbivory on exotics on both continents is not purely a pattern of enemy release in the strictest sense, but to some degree additionally reflects the susceptibility of native plants to invasive herbivores. In conclusion, our cross-Atlantic study is consistent with the idea that nonnative trees have generally reduced herbivory, regardless of tree community diversity and species identity, but for different reasons depending on the dominant herbivore guild.

Quantitative Prediction of Interactions in Bipartite Networks Based on Traits, Abundances, and Phylogeny.

AbstractEcological interactions link species in networks. Loss of species from or introduction of new species into an existing network may have substantial effects for interaction patterns. Predicting changes in interaction frequency while allowing for rewiring of existing interactions-and hence estimating the consequences of community compositional changes-is thus a central challenge for network ecology. Interactions between species groups, such as pollinators and flowers or parasitoids and hosts, are moderated by matching morphological traits or sensory clues, most of which are unknown to us. If these traits are phylogenetically conserved, however, we can use phylogenetic distances to construct latent, surrogate traits and try to match those across groups, in addition to observed traits. Understanding how important traits and trait matching are, relative to abundances and chance, is crucial to estimating the fundamental predictability of network interactions. Here, we present a statistically sound approach ("tapnet") to fitting abundances, traits, and phylogeny to observed network data to predict interaction frequencies. We thereby expand existing approaches to quantitative bipartite networks, which so far have failed to correctly represent the nonindependence of network interactions. Furthermore, we use simulations and cross validation on independent data to evaluate the predictive power of the fit. Our results show that tapnet is on a par with abundance-only, matching centrality, and machine learning approaches. This approach also allows us to evaluate how well current concepts of trait matching work. On the basis of our results, we expect that interactions in well-sampled networks can be well predicted if traits and abundances are the main driver of interaction frequency.

Reply to: "Research on agroforestry systems and biodiversity conservation: what can we conclude so far and what should we improve?" by Boinot et al. 2022.

In our article 'European agroforestry has no unequivocal effect on biodiversity: a time-cumulative meta-analysis' (BMC Ecology and Evolution, 2021) we synthesize the effect of agroforestry on biodiversity. Boinot et al. (BMC Ecology and Evolution, 2022) criticise our approach arguing that our definitions of agroforestry and biodiversity are too narrow; that we use inappropriate control sites for primary studies lacking distance to the treatment sites; that there are too few studies for a meta-analysis in silvoarable systems; and that local practice should be emphasized. We agree on the importance of local practices and that more studies would improve the robustness of our meta-analytical results. However, our conclusions are robust to removing studies criticised for inappropriate controls. We also recognize the problem of different definitions of agroforestry and using species richness as sole proxy for biodiversity. We appreciate being given the opportunity to clarify our results and to trigger future discussions about definitions and the interpretation of results from meta-analysis.

European agroforestry has no unequivocal effect on biodiversity: a time-cumulative meta-analysis.

BACKGROUND: Agroforestry is a production system combining trees with crops or livestock. It has the potential to increase biodiversity in relation to single-use systems, such as pastures or cropland, by providing a higher habitat heterogeneity. In a literature review and subsequent meta-analysis, we investigated the relationship between biodiversity and agroforestry and critically appraised the underlying evidence of the results. RESULTS: Overall, there was no benefit of agroforestry to biodiversity. A time-cumulative meta-analysis demonstrated the robustness of this result between 1991 and 2019. In a more nuanced view silvopastoral systems were not more diverse in relation to forests, pastures or abandoned silvopastures. However, silvoarable systems increased biodiversity compared to cropland by 60%. A subgroup analysis showed that bird and arthropod diversity increased in agroforestry systems, while bats, plants and fungi did not. CONCLUSION: Agroforestry increases biodiversity only in silvoarable systems in relation to cropland. But even this result is of small magnitude, and single-study effect sizes were heterogeneous with sometimes opposing conclusions. The heterogeneity suggests the importance of other, usually unmeasured variables, such as landscape parameters or land-use history, influencing biodiversity in agroforestry systems.

Within-day dynamics of plant-pollinator networks are dominated by early flower closure: an experimental test of network plasticity.

Temporal variability of plant-pollinator interactions is important for fully understanding the structure, function, and stability of plant-pollinator networks, but most network studies so far have ignored within-day dynamics. Strong diel dynamics (e.g., a regular daily cycle) were found for networks with Cichorieae, which typically close their flowers around noon. Here, we experimentally prevented early flower closure to test whether these dynamics are driven by the temporally limited availability of Cichorieae, or by timing of pollinator activity. We further tested if the dynamics involving Cichorieae and their pollinators also affect the dynamics on other plants in the network. Finally, we explored the structure of such manipulated networks (with Cichorieae available in the morning and afternoon) compared to unmanipulated controls (Cichorieae available only in the morning). We found that flower closure of Cichorieae is indeed an important driver of diel network dynamics, while other drivers of pollinator timing appeared less important. If Cichorieae flowers were available in the afternoon, they were visited by generalist and specialist pollinators, which overall decreased link turnover between morning and afternoon. Effects of afternoon availability of Cichorieae on other plants in the network were inconclusive: pollinator switching to and from Cichorieae tended to increase. On the level of the aggregated (full-day) network, the treatment resulted in increased dominance of Cichorieae, reducing modularity and increasing plant generality. These results highlight that network dynamics can be predicted by knowledge of diel or seasonal phenology, and that fixed species timing assumptions will misrepresent the expected dynamics.

Seeing through the static: the temporal dimension of plant-animal mutualistic interactions.

Most studies of plant-animal mutualistic networks have come from a temporally static perspective. This approach has revealed general patterns in network structure, but limits our ability to understand the ecological and evolutionary processes that shape these networks and to predict the consequences of natural and human-driven disturbance on species interactions. We review the growing literature on temporal dynamics of plant-animal mutualistic networks including pollination, seed dispersal and ant defence mutualisms. We then discuss potential mechanisms underlying such variation in interactions, ranging from behavioural and physiological processes at the finest temporal scales to ecological and evolutionary processes at the broadest. We find that at the finest temporal scales (days, weeks, months) mutualistic interactions are highly dynamic, with considerable variation in network structure. At intermediate scales (years, decades), networks still exhibit high levels of temporal variation, but such variation appears to influence network properties only weakly. At the broadest temporal scales (many decades, centuries and beyond), continued shifts in interactions appear to reshape network structure, leading to dramatic community changes, including loss of species and function. Our review highlights the importance of considering the temporal dimension for understanding the ecology and evolution of complex webs of mutualistic interactions.

Increasing connectivity enhances habitat specialists but simplifies plant-insect food webs.

Strong declines of grassland species diversity in small and isolated grassland patches have been observed at local and landscape scales. Here, we study how plant-herbivore interaction webs and habitat specialisation of leafhopper communities change with the size of calcareous grassland fragments and landscape connectivity. We surveyed leafhoppers and plants on 14 small (0.1-0.6 ha) and 14 large (1.2-8.8 ha) semi-natural calcareous grassland fragments in Central Germany, differing in isolation from other calcareous grasslands and in the percentage of arable land in the surrounding landscape (from simple to complex landscapes). We quantified weighted trophic links between plants and their phytophagous leafhoppers for each grassland fragment. We found that large and well-connected grassland fragments harboured a high portion of specialist leafhopper species, which in turn yielded low interaction diversity and simple plant-leafhopper food webs. In contrast, small and well-connected fragments exhibited high levels of generalism, leading to higher interaction diversity. In conclusion, food web complexity appeared to be a poor indicator for the management of insect diversity, as it is driven by specialist species, which require high connectivity of large fragments in complex landscapes. We conclude that habitat specialists should be prioritized since generalist species associated with small fragments are also widespread in the surrounding landscape matrix.

Plant species richness increases with light availability, but not variability, in temperate forests understorey.

BACKGROUND: Temperate forest understorey vegetation poses an excellent study system to investigate whether increases in resource availability lead to an increase in plant species richness. Most sunlight is absorbed by the species-poor tree canopy, making the much more species-rich understorey species inhabit a severely resource-limited habitat. Additionally, the heterogeneity of light availability, resulting from management-moderated tree composition and age structure, may contribute to species coexistence. One would therefore expect that the diversity in the herb layer correlates positively with either the overall light availability, or the light heterogeneity, depending on whether resource availability or heterogeneity are more important drivers of diversity. To test this idea, we assessed variability of light conditions in 75 forest plots across three ecoregions with four different methods. RESULTS: We correlated these data with vegetation relevés and found light availability to be strongly positively correlated with understorey plant species richness, as well as with understorey cover. Light variability (assessed with two approaches) within plots was positively correlated with transmittance, but did not improve the relationship further, suggesting that the main driver of species richness in this system is the overall resource availability. Two of the three beech-dominated regions exhibited near-identical effects of light transmittance, while the third, featuring pine alongside beech and thus with the longest gradient of transmittance and lowest species richness, displayed a weaker light response. CONCLUSIONS: While site conditions are certainly responsible for the trees selected by foresters, for the resulting forest structure, and for the differences in plant species pools, our results suggest that light transmittance is a strong mediating factor of understorey plant species richness.

Co-occurrence patterns and the large-scale spatial structure of benthic communities in seagrass meadows and bare sand.

BACKGROUND: Species distribution models are commonly used tools to describe diversity patterns and support conservation measures. There is a wide range of approaches to developing SDMs, each highlighting different characteristics of both the data and the ecology of the species or assemblages represented by the data. Yet, signals of species co-occurrences in community data are usually ignored, due to the assumption that such structuring roles of species co-occurrences are limited to small spatial scales and require experimental studies to be detected. Here, our aim is to explore associations among marine sandy-bottom sediment inhabitants and test for the structuring effect of seagrass on co-occurrences among these species across a New Zealand intertidal sandflat, using a joint species distribution model (JSDM). RESULTS: We ran a JSDM on a total of 27 macrobenthic species co-occurring in 300,000 m2 of sandflat. These species represented all major taxonomic groups, i.e. polychaetes, bivalves and crustaceans, collected in 400 sampling locations. A number of significant co-occurrences due to shared habitat preferences were present in vegetated areas, where negative and positive correlations were approximately equally common. A few species, among them the gastropods Cominella glandiformis and Notoacmea scapha, co-occurred randomly with other seagrass benthic inhabitants. Residual correlations were less apparent and mostly positive. In bare sand flats shared habitat preferences resulted in many significant co-occurrences of benthic species. Moreover, many negative and positive residual patterns between benthic species remained after accounting for habitat preferences. Some species occurring in both habitats showed similarities in their correlations, such as the polychaete Aglaophamus macroura, which shared habitat preferences with many other benthic species in both habitats, yet no residual correlations remained in either habitat. CONCLUSIONS: Firstly, analyses based on a latent variable approach to joint distributions stressed the structuring role of species co-occurrences beyond experimental scales. Secondly, results showed context dependent interactions, highlighted by species having more interconnected networks in New Zealand bare sediment sandflats than in seagrass meadows. These findings stress the critical importance of natural history to modelling, as well as incorporating ecological reality in SDMs.

Spatial conservation prioritisation in data-poor countries: a quantitative sensitivity analysis using multiple taxa.

BACKGROUND: Spatial conservation prioritisation (SCP) is a set of computational tools designed to support the efficient spatial allocation of priority areas for conservation actions, but it is subject to many sources of uncertainty which should be accounted for during the prioritisation process. We quantified the sensitivity of an SCP application (using software Zonation) to possible sources of uncertainty in data-poor situations, including the use of different surrogate options; correction for sampling bias; how to integrate connectivity; the choice of species distribution modelling (SDM) algorithm; how cells are removed from the landscape; and two methods of assigning weights to species (red-list status or prediction uncertainty). Further, we evaluated the effectiveness of the Egyptian protected areas for conservation, and spatially allocated the top priority sites for further on-the-ground evaluation as potential areas for protected areas expansion. RESULTS: Focal taxon (butterflies, reptiles, and mammals), sampling bias, connectivity and the choice of SDM algorithm were the most sensitive parameters; collectively these reflect data quality issues. In contrast, cell removal rule and species weights contributed much less to overall variability. Using currently available species data, we found the current effectiveness of Egypt's protected areas for conserving fauna was low. CONCLUSIONS: For SCP to be useful, there is a lower limit on data quality, requiring data-poor countries to improve sampling strategies and data quality to obtain unbiased data for as many taxa as possible. Since our sensitivity analysis may not generalise, conservation planners should use sensitivity analyses more routinely, particularly relying on more than one combination of SDM algorithm and surrogate group, consider correction for sampling bias, and compare the spatial patterns of predicted priority sites using a variety of settings. The sensitivity of SCP to connectivity parameters means that the responses of each species to habitat loss are important knowledge gaps.

Exploration of Concerns about the Evidence-Based Guideline Approach in Conservation Management: Hints from Medical Practice.

The importance of using evidence in decision-making is frequently highlighted in policy reports and scientific papers. However, subjective judgments of the reliability of environmental evidence vary widely, and large-scale systematic searches for evidence are only common for climate-related topics. In the medical field, evidence-based guidelines are routinely used to guide treatments. In the management of multiple-use landscapes similar guidelines could substantially narrow the science-practice gap but are largely absent. The challenges potential guidelines face are therefore unknown. For the case of forest conservation, we conducted 14 semistructured interviews with mainly forest practitioners and presented them an example medical guideline together with evidence-based statements on forest conservation (hereinafter: statement paper). We identified 28 concerns related to potential evidence-based guidelines in forest conservation. The interviews yielded approximately three major findings. First, recommendations on forest conservation are better accepted if they include clear instructions and are formulated for a specific context. Fragmentary conservation evidence complicates the formulation of specific recommendations. Second, the level of evidence framework, which indicates the strength of the available evidence, is perceived as too complex. Third, neglecting forest multifunctionality in a potential guideline hampers its application but, if addressed, potentially weakens its ecological relevance. We show that major concerns about potential evidence-based conservation guidelines are similar to the challenges experienced by medical guidelines. We also identify concerns unique to forestry.

Evaluating the effectiveness of retention forestry to enhance biodiversity in production forests of Central Europe using an interdisciplinary, multi-scale approach.

Retention forestry, which retains a portion of the original stand at the time of harvesting to maintain continuity of structural and compositional diversity, has been originally developed to mitigate the impacts of clear-cutting. Retention of habitat trees and deadwood has since become common practice also in continuous-cover forests of Central Europe. While the use of retention in these forests is plausible, the evidence base for its application is lacking, trade-offs have not been quantified, it is not clear what support it receives from forest owners and other stakeholders and how it is best integrated into forest management practices. The Research Training Group ConFoBi (Conservation of Forest Biodiversity in Multiple-use Landscapes of Central Europe) focusses on the effectiveness of retention forestry, combining ecological studies on forest biodiversity with social and economic studies of biodiversity conservation across multiple spatial scales. The aim of ConFoBi is to assess whether and how structural retention measures are appropriate for the conservation of forest biodiversity in uneven-aged and selectively harvested continuous-cover forests of temperate Europe. The study design is based on a pool of 135 plots (1 ha) distributed along gradients of forest connectivity and structure. The main objectives are (a) to investigate the effects of structural elements and landscape context on multiple taxa, including different trophic and functional groups, to evaluate the effectiveness of retention practices for biodiversity conservation; (b) to analyze how forest biodiversity conservation is perceived and practiced, and what costs and benefits it creates; and (c) to identify how biodiversity conservation can be effectively integrated in multi-functional forest management. ConFoBi will quantify retention levels required across the landscape, as well as the socio-economic prerequisites for their implementation by forest owners and managers. ConFoBi's research results will provide an evidence base for integrating biodiversity conservation into forest management in temperate forests.

Adaptation of wild boar (Sus scrofa) activity in a human-dominated landscape.

BACKGROUND: Wild boars (Sus scrofa L.) are globally widely distributed, and their populations have increased in Europe during recent decades. Encounters between humans and wild boars are rare because of the predominantly nocturnal lifestyle of the latter, and wild boar management by hunting is a challenging task. Animal activity patterns are important for understanding the behaviour of a species. However, knowledge of detailed temporal patterns and an understanding of the drivers of wild boar activity at a fine temporal scale are lacking. Of special relevance for human-wild boar interactions (e.g., encounters, conflicts, and management) is the question of whether nocturnal activity depends on anthropogenic factors and, particularly, how local hunting regimes may affect activity patterns. We used GPS telemetry and acceleration measurements to shed light on this part of wild boar behaviour, observing 34 animals in Central Europe. Animals were tracked along a gradient of hunting pressure from hunting-free areas to areas with low or high hunting pressure. Fitted generalised additive models allowed predicting the probability of active behaviour under differing disturbance regimes precisely to day of year and time of day. RESULTS: The wild boars were predominantly nocturnal, with peak activity at approximately midnight. However, the data showed increased activity during daylight for wild boars that used no-hunting zones or reduced-hunting zones. Large areas with low disturbance levels promoted activity during daylight more than smaller areas with an intermediate disturbance regime. High air temperatures and locations within forests reduced the probability of active behaviour, whereas proximity to tracks used for forestry or agriculture was accompanied by a higher probability of activity. CONCLUSIONS: We conclude that wild boars flexibly adjust their activity to their local environmental conditions, considering disturbances at the scale of long-term home ranges as well as actual small-scale landscape quality. Entire wild boar home ranges should be covered in the delineation of reserves intending to stimulate activity during daylight.

A new model explaining the origin of different topologies in interaction networks.

Nestedness and modularity have been recurrently observed in species interaction networks. Some studies argue that those topologies result from selection against unstable networks, and others propose that they likely emerge from processes driving the interactions between pairs of species. Here we present a model that simulates the evolution of consumer species using resource species following simple rules derived from the integrative hypothesis of specialization (IHS). Without any selection on stability, our model reproduced all commonly observed network topologies. Our simulations demonstrate that resource heterogeneity drives network topology. On the one hand, systems containing only homogeneous resources form generalized nested networks, in which generalist consumers have higher performance on each resource than specialists. On the other hand, heterogeneous systems tend to have a compound topology: modular with internally nested modules, in which generalists that divide their interactions between modules have low performance. Our results demonstrate that all real-world topologies likely emerge through processes driving interactions between pairs of species. Additionally, our simulations suggest that networks containing similar species differ from heterogeneous networks and that modules may not present the topology of entire networks.

Refuges from fire maintain pollinator-plant interaction networks.

Fire is a major disturbance factor in many terrestrial ecosystems, leading to landscape transformation in fire-prone areas. Species in mutualistic interactions are often highly sensitive to disturbances like fire events, but the degree and complexity of their responses are unclear. We use bipartite insect-flower interaction networks across a recently burned landscape to explore how plant-pollinator interaction networks respond to a recent major fire event at the landscape level, and where fire refuges were present. We also investigate the effectiveness of these refuges at different elevations (valley to hilltop) for the conservation of displaced flower-visiting insects during fire events. Then, we explore how the degree of specialization of flower-visiting insects changes across habitats with different levels of fire impact. We did this in natural areas in the Greater Cape Floristic Region (GCFR) biodiversity hotspot, which is species rich in plants and pollinators. Bees and beetles were the most frequent pollinators in interactions, followed by wasps and flies. Highest interaction activity was in the fire refuges and least in burned areas. Interactions also tracked flower abundance, which was highest in fire refuges in the valley and lowest in burned areas. Interactions consisted mostly of specialized flower visitors, especially in refuge areas. The interaction network and species specialization were lowest in burned areas. However, species common to at least two fire classes showed no significant difference in species specialization. We conclude that flower-rich fire refuges sustain plant-pollinator interactions, especially those involving specialized species, in fire-disturbed landscape. This may be an important shelter for specialized pollinator species at the time that the burned landscape goes through regrowth and succession as part of ecosystem recovery process after a major fire event.

African elephant poaching rates correlate with local poverty, national corruption and global ivory price.

Poaching is contributing to rapid declines in elephant populations across Africa. Following high-profile changes in the political environment, the overall number of illegally killed elephants in Africa seems to be falling, but to evaluate potential conservation interventions we must understand the processes driving poaching rates at local and global scales. Here we show that annual poaching rates in 53 sites strongly correlate with proxies of ivory demand in the main Chinese markets, whereas between-country and between-site variation is strongly associated with indicators of corruption and poverty. Our analysis reveals a recent decline in annual poaching mortality rate from an estimated peak of over 10% in 2011 to <4% in 2017. Based on these findings, we suggest that continued investment in law enforcement could further reduce poaching, but is unlikely to succeed without action that simultaneously reduces ivory demand and tackles corruption and poverty.

Spatial behavior in rehabilitated orangutans in Sumatra: Where do they go?

Wildlife restoration is one of the key components of conservation strategies, and this includes the rehabilitation and release of animals confiscated from wildlife traffickers. When primates are re-introduced, most individuals need a pre-release training to acquire the skills needed to survive in the wild. Pre-release training may either negatively or positively affect primate post-release behavior and survival. Post-release behavior, however, has rarely been monitored even though it is the only means to assess the survival of released individuals. Here, we present a thorough analysis of data from a 3-year radio tracking study on 32 orangutans (Pongo abelii) released in Sumatra after their rehabilitation. We investigated whether and how the age at release, the duration of the pre-release rehabilitation and training, and the release location affected the post-release individual spatial behavior. Orangutans released at older age exhibited post-release habitat selection patterns that were more comparable to that shown by wild conspecifics, i.e., they chose areas closer to rivers and at lower elevations (150-250 meters a.s.l.) where previous research had documented greater food availability. In contrast, individuals released at younger age showed a stronger spatial dependency on the rehabilitation station and exhibited disrupted habitat selection patterns; although after several months after the release all individuals tended to decrease their spatial reliance on the rehabilitation facility. This study indicates that the rehabilitation of individuals for a longer period and their release further from the rehabilitation station have facilitated the subsequent development of more natural spatial behavior, i.e. driven by food availability rather than by the dependence on care-giving human facility. Our study provides indications on how to improve the rehabilitation and release of confiscated orangutans, highlighting the importance of the age at release, the length of the rehabilitation program, and the location of the release site.