Humans have inhabited Bornean tropical forests for at least 45,000 years, yet the impact of that long-term presence is poorly understood. Borneo's extensive archaeological record and high biodiversity offer a unique laboratory to examine patterns of site selection and the effects of human activity on tropical forest ecosystems. This paper establishes a synthetic literature-based data set of 73 archaeological sites, 47 with radiocarbon or radiometric records dating during the last 6 ka BP. Palaeoclimate, current climate, soil, and terrain variables of these 47 sites are compared against a set of simulated random locations across Borneo. We also consider spatial proximity as a factor influencing human settlement, clustering sites based on location and examining the timing and duration of habitation of sites within the clusters. Finally, we compare the 47 site records against the HYDE-3.2 land-use model to assess correspondence. Our results suggest that during the last 6000 years BP, Bornean human settlements tended to be located in the forests at lower elevations, near coasts and river networks, in higher temperatures and lower precipitation. Comparative sedimentary analysis also suggests preferential use of locations exhibiting lower clay and higher sand content. This combination of conditions likely improved food production and resource supply. Our study highlights the value of integrating archaeological data with global historical land-use and climate models to uncover long-term human-environment interactions. By establishing a cross-site environmental baseline, these findings provide insights into past human settlement patterns and likely human legacies in Borneo's tropical forests.

Parasites can have large impacts on host populations, but the extent to which parasite dynamics impact or respond to multi-species community structure remains uncertain. Empirical and theoretical studies within the host-microbiome feedback framework (often called plant-soil feedback) has provided strong evidence of the importance of soil pathogens to plant community structure and function. We adapt this framework to herd animals by extending the mathematics of host-microbiome feedback theory to accommodate increased likelihood of exposure to microbiomes from conspecific hosts rather than heterospecific hosts. We then integrate this framework with a model of interguild frequency dependence. Coupling this model with empirical observations, we estimate the host-specific fitness of gastro-intestinal nematodes living on ungulate species of Western USA. We find evidence that host-specific differences in nematode fitness could generate negative feedback on host fitness and contribute to coexistence of ungulates. Moreover, we find that this is more likely to be the case for pairs of ungulate species with high habitat overlap. If nematodes can indeed drive such negative feedbacks, then negative impacts of nematodes on their ungulate hosts should decline, i.e., be diluted, with increasing host diversity. While more work is necessary to confirm the underlying assumptions driving these conclusions, our work highlights the possibility that parasites play under appreciated roles in structuring animal communities.

The rapid environmental changes of the Anthropocene create legacy effects that may shape future Earth system responses. One significant legacy effect is the species extinction debt caused by past habitat destruction. As biodiversity underpins ecosystem services vital to human societies, social-ecological systems may, in turn, be subjected to biodiversity-dependent ecosystem service debts. While biodiversity-dependent ecosystem service debts have been quantified with analytical approaches, less attention has been paid to their potential impact on social-ecological system trajectories. We performed a theoretical study of a dynamical systems model that includes the possibility of biodiversity-dependent ecosystem service debts emerging from past habitat destruction. Our results suggest that these debts reduce systems' safe operating spaces and create environmental tipping points associated with critical transitions in system states. These transitions, however, may include long transients of apparent stability, making it difficult to identify cause and effect. Notably, biodiversity-dependent ecosystem service debts may drive initial phases of apparent recovery after disturbance, still followed by system collapse. Our theoretical findings highlight the need to consider biodiversity-dependent ecosystem service debts for sustainable management of social-ecological systems. Furthermore, these results suggest that social-ecological systems' safe operating spaces cannot be reliably inferred from recent observations of apparent system stability.

Fires are generally considered to promote biodiversity, although this relationship can be affected by several factors, including fire regime and ecosystem type. Given the ongoing global change, a better understanding of this connection is needed to assess the extent to which projected increases in fire frequency may affect current biodiversity trends. A major challenge lies in vegetation-fire feedback, which often mediates the changes in fire regime. To shed light on the role of fires in promoting or limiting biodiversity, we studied the compositional and functional diversity of simulated plant communities as a function of different fire frequencies. We extended an existing model to include a large number of species. The model reproduces successional dynamics and is parameterized to represent Boreal and Mediterranean plant communities. Fire events are stochastic, with frequency that depends on the community flammability, and plants have different fire responses, thus representing a vegetation-fire feedback. For both ecosystems, we found that fires generally have a positive effect on diversity. All of the biodiversity indicators analyzed, to some extent, peaked at intermediate fire frequency. Interestingly, we observe a decoupling between compositional and functional diversity. These results are related to the vegetation-fire feedback and underline its importance for biodiversity in fire-prone ecosystems. ### Competing Interest Statement The authors have declared no competing interest. Piano Nazionale di Ripresa e Resilienza PNRR, P2022NRLF2 Italian National Biodiversity Future Center, CN 00000033 European Union-NextGenerationEU

Sustainability and environmental impact assessments of trade and investment agreements need to address biodiversity more effectively. To showcase this, we examined a report, commissioned by Switzerland, on potential environmental impacts and risks of the trade between Switzerland and the Mercosur under the EFTA–Mercosur Free Trade Agreement. Our analysis focuses on chemical pollution, regulatory impact, duty-free precious minerals, deforestation, and greenhouse gas emission to develop a roadmap for (re-)interpreting (muted) risks on biodiversity in light of the Global Biodiversity Framework. Our analytical approach on normative and substantive effectiveness of impact assessments encourages broader responses to impacts of economic policies on biodiversity and ecosystems.

Land-cover transitions from forest to non-forest land-covers have led to substantial loss and fragmentation of global forests. While average rates of forest change are well studied, less attention has been paid to spatio-temporal trajectories of forest loss and recovery. These trajectories likely differ by land-cover transition types, and may be constrained by the initial amount and spatial arrangement of the non-forest land-cover. In this study, we distinguish between abrupt and gradual forest change trajectories, evaluating how they vary across land-cover transitions and how initial non-forest land-cover conditions mediate the amount and rate of forest change. To this end, we use annual global land-cover maps (1992–2020) from the European Space Agency’s Climate Change Initiative. We find that the expansion of croplands and bare lands drive forest loss, while transitions from wetlands, shrublands and grasslands are key contributors to forest gain. Forest loss rates were lower when the non-forest land-cover was initially fragmented, except for settlements and bare land, where fragmentation accelerated loss. Conversely, forest gain was highest in wetlands and shrublands, though increased fragmentation generally reduced the amount and rate of forest gain. Our results show that the amount and rate of forest change are associated with the initial amount and spatial arrangement of the non-forest, transition land-cover. This insight may improve our ability to link remotely-sensed land-cover changes to the underlying drivers of global forest change.

The speed and severity of tropical forest conversions to croplands and rangelands, give rise to distinct fragmentation patterns, altering species composition and ecosystem resilience. While the type of the surrounding land-cover can mitigate these impacts through landscape complementation, the role of specific land-cover transitions and their speeds in shaping forest fragmentation patterns remains poorly understood at the pan-tropical scale. We study how fragmentation dynamics vary by the type, speed and severity of the land-cover transition, while also accounting for the constraints imposed by the initial forest amount. We hypothesize that forest fragmentation patterns will vary by transition type, with rapid transitions producing compact forest patches with distinct geometric boundaries. We analyzed fragmentation dynamics across 87,510 tropical forest landscapes of size 30.09 km2 using global land-cover maps from 1992–2020. Our results show that: (1) grasslands exhibit a patchy and dispersed pattern of expansion, while croplands and shrublands exhibit threshold behavior—triggering rapid fragmentation once initial forest cover falls below 60

Purpose The GreenComp framework identifies 12 competences for sustainability as common ground for higher-education curricula. The framework can be used for self-assessment and the review of curricula. However, a step-by-step method to conduct such a self-assessment is not yet available for the GreenComp framework specifically. Therefore, the authors present the GreenComp Evaluation Roadmap allowing to evaluate the extent to which the frameworks’ competences for sustainability are integrated in higher-education curricula. The application of the GreenComp Evaluation Roadmap to a curriculum taught at the University of Aruba, allows to report on the benefits, limitations and future potential of the approach. Design/methodology/approach The proposed mixed-method approach combines hybrid qualitative and quantitative data collection on the integration of the 12 competences for sustainability of the GreenComp framework in higher-education curricula. The authors showcase its potential through application of the GreenComp framework as an evaluation tool to a science, technology, engineering and mathematics-based bachelor program taught at the University of Aruba. Findings The GreenComp Evaluation Roadmap not only allows for an evaluation of the curriculum and identification of competence gaps. It also supports educators to conduct a self-reflection on individual course(s) and the program as a whole. The paper shows promising results that the roadmap developed could be a reproducible approach. Moreover, it provides guidance to other higher education institutes for self-evaluation and self-reflection on how the competences for sustainability are integrated in their curricula and how this can be enhanced in the future. Originality/value The need to integrate sustainability throughout higher-education curricula is broadly recognized. The GreenComp Evaluation Roadmap contributes to the literature by offering a methodological approach to evaluate the integration of the 12 competences for sustainability throughout a curriculum.

Global biodiversity loss and climate change exacerbate feedbacks within social-ecological systems, i.e., between ecosystems, their services and well-being of human societies. Our ability to mediate these feedbacks is hampered by incomplete understanding of the underlying causal links, which could benefit from interdisciplinary approaches to discover theoretical or empirical links from heterogeneous data characteristic of social-ecological studies. We propose a novel framework connecting literature-based causal knowledge with data-driven inference of causality. We test this framework for the highly biodiverse island of Borneo by conducting a systematic literature review of 7473 studies over 170 years, and a causal inference analysis for three conceptual causal diagrams connecting global change, socio-economics, ecosystem services, and biodiversity-ecosystem function using a set of 227 spatially explicit variables. We find that, while natural or social processes have been mostly studied independently, a set of studies already documents causal links across social-ecological domains for processes related to deforestation, food or energy. Causal discovery unveiled consistent negative causal links between global change, social-economic landscape, and biodiversity-ecosystem function, and positive causal links between global change and socio-economics, and these links were robust to indicator selection and addition. We detected few and weak links between social-economic landscape, global change, and ecosystem services. When comparing the data-driven inferred causal links to those documented by the literature, we find that links between biodiversity and ecosystem function with global change, and links between social-economic landscape and ecosystem services were also consistent, and causal analysis uncovered new (potential) causal links not yet described in the literature. Significance Statement Addressing climate change and biodiversity loss in the Anthropocene requires us to recognize that human societies and ecological systems are inherently interconnected in complex adaptive systems. Causal understanding in social-ecological systems enables understanding system dynamics and response to pressures and shocks. While promising, few studies have studied these systems using a combination of ‘big literature’ which provides the state-of-the-knowledge and ‘big data’ that provides the underlying information for causal discovery. With this framework, we can specify and rigorously test, causal links in biodiversity-mediated social-ecological processes under global change and examine potential interventions that lead to much needed sustainable outcomes. ### Competing Interest Statement The authors have declared no competing interest.

PurposeUniversities of Small Island States (SIS) have the potential to fulfill a crucial role in implementing the Sustainable Development Goals (SDGs) but also face barriers to local capacity building. The University of Aruba partly addresses these challenges through the development of The Academic Foundation Year (AFY), a one-year pre-university program aiming to optimally equip students for higher education. This study aims to assess to what extent the program can foster the local embeddedness of the students in ecology, culture and history and an understanding of opportunities and challenges for sustainable development in SIS. Design/methodology/approachThe authors provide examples of how the program incorporates education for sustainable development and how it emphasizes experiential learning. In addition, quantitative survey data and qualitative analysis of focus group meetings are used to reflect on the program's achievements and its potential for further development. FindingsThe survey results suggest that AFY courses not only increase knowledge but also change students' perceptions regarding sustainability. Indeed, key impacts emerging from the student focus group related to both academic preparation and engagement with sustainability. Reflections by teachers emphasized the importance of experiential learning, an expansive view of the SDGs and preparing students as citizens. Originality/valueThis study highlights that the program could provide a starting point for the development of similar initiatives in other SIS, the common basis being the fostering of sustainability literacy and social adoption of the SDGs.

Navigating social-ecological systems toward sustainable trajectories is an important challenge of the Anthropocene. Models of social-ecological systems can increase our understanding of how social and ecological subsystems interact, their response to environmental changes, and how their dynamics may be altered by management interventions. However, the level of representational detail required for models to describe a particular social-ecological system with high fidelity (i.e., accurately quantifying system dynamics) may hamper both the interpretability of model results and our ability to identify key processes and feedbacks within the system. In contrast, stylized models describe simplified interactions between a small subset of social-ecological system elements. Stylized models are a useful tool to identify potential consequences of specific key processes and feedbacks on system functioning. However, the relatively low level of representational detail in these models limits their ability to deliver concrete management options for a particular social-ecological system. Here, we describe how an exploratory modeling approach can utilize the strengths of stylized models before the construction of social-ecological system models with high fidelity and representational detail. This exploratory modeling approach is an iterative strategy, with the initial steps comprising the development of stylized models informed by empirical observations. We illustrate this with two examples of stylized modeling of isolated and connected social-ecological systems. Through repeated confrontation of alternative models with empirical data, exploratory modeling provides useful stepping stones toward the development of models that describe social-ecological systems in increasingly specific settings with increasing levels of representational detail. When these latter types of models reach a high level of fidelity, they could be used for scenario-based analyses and participatory decision-making processes. At this stage, the conceptual insights previously obtained during the exploratory modeling phase may aid in the interpretation and communication of the outcomes of scenario-based analyses. Hence, exploratory modeling aims to create a synergy between the insights obtained from stylized models and system-specific, high-fidelity models in order to generate a deep understanding of the drivers of social-ecological system dynamics, and how to leverage these drivers to initiate desired changes.

The emergence and maintenance of tree species diversity in tropical forests is commonly attributed to the Janzen-Connell (JC) hypothesis, which states that growth of seedlings is suppressed in the proximity of conspecific adult trees. As a result, a JC distribution due to a density-dependent negative feedback emerges in the form of a (transient) pattern where conspecific seedling density is highest at intermediate distances away from parent trees. Several studies suggest that the required density-dependent feedbacks behind this pattern could result from interactions between trees and soil-borne pathogens. However, negative plant-soil feedback may involve additional mechanisms, including the accumulation of autotoxic compounds generated through tree litter decomposition. An essential task therefore consists in constructing mathematical models incorporating both effects showing the ability to support the emergence of JC distributions.In this work, we develop and analyse a novel reaction-diffusion-ODE model, describing the interactions within tropical tree species across different life stages (seeds, seedlings, and adults) as driven by negative plant-soil feedback. In particular, we show that under strong negative plant-soil feedback travelling wave solutions exist, creating transient distributions of adult trees and seedlings that are in agreement with the Janzen-Connell hypothesis. Moreover, we show that these travelling wave solutions are pulled fronts and a robust feature as they occur over a broad parameter range. Finally, we calculate their linear spreading speed and show its (in)dependence on relevant nondimensional parameters.

By 2050, 70% of the global population will reside in urban areas, raising concerns about food security and biodiversity loss. Tropical homegardens, a form of agroforestry, may have the potential to align concurrent demands for biodiversity conservation and food security in urban environments. Nevertheless, this depends on land-use decision-making strategies that are enabled by household livelihood assets. We examined which livelihood assets are associated with homegarden agrobiodiversity and food availability and accessibility in Kerala, India, the nature of these associations, and whether they vary along urbanisation gradients using data from 304 homegardens. We observed higher plant species richness closer to the urban centre and in smaller homegardens. Meanwhile, yields and share of household food consumption from homegardens tend to be higher farther away from urban centres. Assets like fertilizer use, irrigation and kinship affiliation are positively associated with both homegarden agrobiodiversity and food outcomes. Conversely, homegarden size is negatively associated with biodiversity outcomes across urbanisation gradients and positively associated with yield outcomes furthest away from urban centre. Some assets like family labour and farming association membership were only positively associated with food outcomes, while others like external labour availability were positively associated only with plant diversity in the urban centre. Overall, synergies between homegarden agrobiodiversity and household food security requires inputs to natural processes, such as irrigation and soil fertility, and social processes like knowledge sharing and social cohesion, especially in urban peripheries. We emphasize the relevance of homegardens and enhancing livelihood assets for food-secure households in biodiverse urban environments.

Ecosystem services are often analysed individually, but are intertwined with one another and the social-ecological systems they occur in. As a response, ecosystem service bundles, i.e. co-occurring sets of ecosystem services, can be used to simplify complex relationships between nature and society, and in turn aid understanding. Typically bundles are studied on the local to regional scale, given the importance of local context to bundling, but wider scale analysis may help highlight broader ecosystem service balances for sustainable management. However, it remains uncertain if the relationships between ecosystem services are strong enough to describe coherent bundles at the global scale, and the extent to which these bundles are robust across different social-ecological systems and within different biogeographical realms. Here, we examine whether coherent bundles emerge from a set of 25 ecosystem property and service indicators across regional mountain, island and delta systems around the world. We analyse differences between bundle composition and correlation structure based on system, latitude and biome. We find consistent bundles broadly representing 'food', 'productivity' and biodiversity 'intactness/soil' ecosystem properties and services emerge across mountains, islands and deltas globally. These bundles show strong positive correlations internally, and consistent negative correlations between 'food' services and 'intactness/soil' ecosystem properties across bundles. The bundles weakened at higher latitudes and individual biomes where the division between ecosystem properties and services broke down. In sum, while islands, mountains and deltas are distinct social-ecological systems, we found ecosystem bundles robustly described synergies and trade-offs between ecosystem services across these systems. This suggests that bundling has a role in simplifying wider scale interactions between humans and ecosystem services.

Choosing restoration strategies may depend on ecosystem's stability properties. When degraded ecosystems do not self‐perpetuate, natural regeneration can lead to system recovery, and restoration interventions are often designed to accelerate the natural regeneration process. However, when degraded systems self‐perpetuate, reestablishing functional ecosystems depends on overcoming resistance thresholds that impede recovery. In both scenarios, concentrating restoration efforts in patches of the desired state may enhance ecosystem recovery. Introducing patches of a desired state has been motivated by two frameworks: autocatalytic nucleation and the analogy to nucleation. When restoration depends on overcoming resistance thresholds, autocatalytic nucleation lowers restoration barriers by initiating a local positive feedback mechanism that is only successful when desired patches are introduced above a critical patch size. In contrast, the analogy to nucleation accelerates natural regeneration whereby desired patches interact with landscape scale factors often through directed dispersal. We compare nucleation frameworks, and discuss their applications for restoration practices.

1. Dispersal plays an important role in the survival and spread of plant populations. However, when grown under stress, a plants' architecture and fecundity may be modified, which can compromise seed dispersal. While populations within a species exhibit trait variation in response to their environment, a mechanistic understanding of how trade-offs between dispersal ability and competitive ability vary across species' ranges is currently lacking. 2. Using a common garden design, populations of a widespread weedy species (Cardamine hirsuta) were grown to (1) characterize how traits that influence dispersal differ between populations and (2) identify whether a trade-off between dispersal ability and competitive ability varies among populations in response to intraspecific competition. We measured trait differences between populations and characterized dispersal kernels through exhaustively capturing seeds. 3. In the absence of competition, populations exhibited differences in fecundity and height. Of these traits, fecundity varied more strongly between populations and had a stronger influence on dispersal capacity. 4. The experimental data were then used to parameterize cellular automaton models to explore the consequences of these trait-mediated dispersal responses for long-term population dynamics. These simulations identified intraspecific competitive effects on fecundity as the primary driver of variation in spread and patch density among populations. 5. Within-species variation of dispersal traits occurs among populations across the range of C. hirsuta. In response to conspecific competition, populations also have unequal fecundity and long-distance dispersal (LDD). Model simulations suggest predictions of species spread are highly modified by characterization of metapopulation life-history traits such as high fecundity and maintenance of LDD. These mechanisms creating within-species variation would be important to incorporate in population models of species range expansion and stability.

Productivity benefits from diversity can arise when compatible pathogen hosts are buffered by unrelated neighbors, diluting pathogen impacts. However, the generality of pathogen dilution has been controversial and rarely tested within biodiversity manipulations. Here, we test whether soil pathogen dilution generates diversity- productivity relationships using a field biodiversity-manipulation experiment, greenhouse assays, and feedback modeling. We find that the accumulation of specialist pathogens in monocultures decreases host plant yields and that pathogen dilution predicts plant productivity gains derived from diversity. Pathogen specialization predicts the strength of the negative feedback between plant species in greenhouse assays. These feedbacks significantly predict the overyielding measured in the field the following year. This relationship strengthens when accounting for the expected dilution of pathogens in mixtures. Using a feedback model, we corroborate that pathogen dilution drives overyielding. Combined empirical and theoretical evidence indicate that specialist pathogen dilution generates overyielding and suggests that the risk of losing productivity benefits from diversity may be highest where environmental change decouples plant-microbe interactions.

Sustainable management of social-ecological systems requires an understanding of how anthropogenic climate -and land use change may disrupt interactions between human societies and the ecosystem processes they depend on. In this study, we expand an existing stylized social-ecological system model by explicitly considering how urbanizing societies may become less dependent on local ecosystem functioning. This expansion is motivated by a previously developed conceptual framework suggesting that societies may reside in either a green loop and be strongly dependent on local ecosystem processes, or in a red loop where this dependency is weaker due to im-ports of natural resources from elsewhere. Analyzing the feasibility and stability of local social-ecological system states over a wide range of environmental and socio-economic conditions, we observed dynamics consistent with the notion of green loop-dominated and red loop-dominated societies comprising alternate stable social -ecological states. Based on systems' inherent dependencies on local ecosystem processes, responses to environ-mental change could comprise either transitions between green loop-and red loop-dominated states, or collapse of either of these states. Our quantitative model provides an internally consistent mapping of green loop-and red loop-dominated states, as well as transitions between or collapses of these states, along a gradient of environ-mental conditions.