Ecological Integrity of Hybrid Ecosystems in the Anthropocene: The Impact of Self-Organisation on Function and Sustainability

Ecological Integrity of Hybrid Ecosystems in the Anthropocene: The Impact of Self-Organisation on Function and Sustainability | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL Ecology and Evolution This is a preprint and has not been peer reviewed. Data may be preliminary. 13 August 2025 V1 Latest version Share on Ecological Integrity of Hybrid Ecosystems in the Anthropocene: The Impact of Self-Organisation on Function and Sustainability Authors : Elli Groner 0000-0002-7843-3047 [email protected] , Aviva Peeters , and Moshe Shachak Authors Info & Affiliations https://doi.org/10.22541/au.175508139.98145484/v1 Published Ecology and Evolution Version of record Peer review timeline 356 views 211 downloads Contents Abstract Title Authors names: Elli Groner, Aviva Peeters and Moshe Shachak A short running title of less than 40 characters Abstract and keywords Data Accessibility Statement Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Introduction The Anthropocene is defined by significant human-driven transformations of Earth’s ecosystems, resulting in hybrid systems that merge natural self-organisation with varying degrees of anthropogenic management. These hybrid ecosystems now dominate the terrestrial biosphere, playing a critical role in sustaining biodiversity, ecosystem functions, and human well-being. The Role of Self-Organisation in Hybrid Ecosystem Processes In this paper, we argue that self-organisation, the spontaneous emergence of structure and function through local ecological interactions, is a fundamental driver of ecosystem resilience and integrity in hybrid systems. In this manuscript, we synthesize six core self-organizing processes: spatial pattern formation, population self-regulation, ecological feedbacks, ecosystem engineering, functional food web organisation, and functional redundancy, to show their role in underpinning key ecosystem functions across diverse anthrome categories. The Anthromes Using the anthrome framework, we demonstrate how self-organisation operates within croplands, rangelands, village mosaics, forest-populated mosaics, and seminatural systems, emphasizing the ecological functions these processes support under varying land-use regimes. Framework for Navigating Self-Organisation and Ecological Integrity of Hybrid Ecosystems Finally, we propose a conceptual framework for assessing ecological integrity in hybrid ecosystems based on the identification of active self-organizing processes and the mapping of their functional contributions. This process-based approach offers a practical and scalable tool for monitoring ecosystem health and guiding adaptive management strategies in an era of escalating environmental change. Title page Title Ecological Integrity of Hybrid Ecosystems in the Anthropocene: The Impact of Self-Organisation on Function and Sustainability Authors names: Elli Groner, Aviva Peeters and Moshe Shachak Author’s affiliation DSASC Dead Sea and Arava Science Center, Ramon Masada National Park, Massada, IL 86910 [email protected] BGU Ben-Gurion University of the Negev - Eilat Campus, Eilat Campus 162 Hatmarim Avenue Eilat, IL 8801401 SCE Negev School of Architecture, Shamoon College of Engineering, Architecture 56 Bialik street Beer Sheva, IL 8434231 TerraVision Lab, GIS 6 Nahal Karkash street Midreshet Ben-Gurion, IL 8000049 BGU Ben-Gurion University of the Negev, Ecology Sde Boqer Campus Midreshet Ben-Gurion, South District, IL 84990000 A short running title of less than 40 characters Self-Organisation in Hybrid Ecosystem Abstract and keywords Abstract: Introduction The Anthropocene is defined by significant human-driven transformations of Earth’s ecosystems, resulting in hybrid systems that merge natural self-organisation with varying degrees of anthropogenic management. These hybrid ecosystems now dominate the terrestrial biosphere, playing a critical role in sustaining biodiversity, ecosystem functions, and human well-being. The Role of Self-Organisation in Hybrid Ecosystem Processes In this paper, we argue that self-organisation, the spontaneous emergence of structure and function through local ecological interactions, is a fundamental driver of ecosystem resilience and integrity in hybrid systems. In this manuscript, we synthesize six core self-organizing processes: spatial pattern formation, population self-regulation, ecological feedbacks, ecosystem engineering, functional food web organisation, and functional redundancy, to show their role in underpinning key ecosystem functions across diverse anthrome categories. The Anthromes Using the anthrome framework, we demonstrate how self-organisation operates within croplands, rangelands, village mosaics, forest-populated mosaics, and seminatural systems, emphasizing the ecological functions these processes support under varying land-use regimes. Framework for Navigating Self-Organisation and Ecological Integrity of Hybrid Ecosystems Finally, we propose a conceptual framework for assessing ecological integrity in hybrid ecosystems based on the identification of active self-organizing processes and the mapping of their functional contributions. This process-based approach offers a practical and scalable tool for monitoring ecosystem health and guiding adaptive management strategies in an era of escalating environmental change. Key words: Anthrome, Anthropocene, Ecosystem, Pattern formation, Stability, Conservation Main text Introduction: The Anthropocene is an epoch marked by profound and pervasive human impacts on Earth’s ecosystems. It features a complex interplay between natural processes and human activities that shape biodiversity and ecosystem functionality (Crutzen, 2002; Steffen et al., 2007). This era is characterized by rapid climate change, biodiversity loss, extensive land-use modifications, and disruptions to global biogeochemical cycles. These transformations significantly influence ecosystem structure, function, and the provision of essential services, including clean air, water, food security, and climate regulation (Steffen et al., 2004). Currently, most of Earth’s terrestrial landscapes can be described as hybrid ecosystems shaped by ecological self-organisation and varying degrees of anthropogenic management. These ecosystems exist along a continuum between naturally self-organized systems, primarily governed by intrinsic ecological processes, and ecosystems predominantly influenced and organized by human intervention (Ellis and Ramankutty 2008). Globally, hybrid ecosystems now account for more than 75% of the terrestrial biosphere, while only 22% to 25% remain minimally affected wildlands (Ellis and Ramankutty 2008; Kennedy et al., 2019). The distribution of hybrid ecosystems varies regionally: densely populated cropland-village mosaics dominate South and Southeast Asia, extensive rangelands and agro-pastoral systems prevail across Sub-Saharan Africa, multifunctional agricultural-residential mosaics are widespread in Europe and North America, and agroforestry landscapes are common in large parts of Latin America and tropical forest frontiers. Indigenous fire and grazing management systems have created resilient hybrid landscapes in Oceania and Australia. Hybrid ecosystems exemplify the dynamic interaction between self-organized ecological processes such as feedback loops, succession, and spatial pattern formation and human-driven modifications, including land-use change, resource extraction, and infrastructure development. This complex interplay necessitates a deeper understanding of the role self-organisation plays in the functioning, sustainability, and ecological integrity of hybrid ecosystems in the Anthropocene. This paper argues that self-organisation plays a critical role in sustaining ecosystem functions and services in hybrid ecosystems, and that assessing the degree and nature of self-organisation can provide a meaningful indicator of their ecological integrity (EI). An ecosystem that is strongly influenced by human activities yet still governed by natural dynamics should be classified as a hybrid ecosystem . In such systems, the relative influence of bottom-up (self-organizing) versus top-down (externally imposed) forces vary. Rather than merely labelling ecosystems as hybrid, it is essential to quantify the relative weight of these forces to better characterize their nature. Some hybrid ecosystems are predominantly self-organized, with anthropogenic influences largely masked by internal ecological dynamics. Others are primarily shaped by external human forces, with self-organizing processes being suppressed or obscured. Still, some systems represent a more balanced interaction between the two. This perspective suggests a continuum —ranging from ecosystems driven largely by self-organisation to those dominated by anthropogenic control. To evaluate the role of self-organisation in the ecological integrity of hybrid ecosystems systematically, we adopted the “ anthrome ” framework developed by Ellis (Ellis & Ramankutty, 2008). This framework classifies human-influenced ecosystems based on population density, land use, and ecological structure. Each anthrome represents a unique configuration of human-nature interactions, combining human land use with varying levels of natural ecosystem structure and self-regulation. These ecosystems are ”hybrids” not only in terms of land use but also in their internal ecological dynamics. Even under intensive human management, self-organizing processes, such as spatial pattern formation, species self-regulation, feedback loops, and ecosystem engineering, continue to operate and affect ecosystem resilience and functionality (Rietkerk et al., 2004; Murdoch, 1994; Jones et al., 1994). Understanding the role of self-organisation is essential for assessing the sustainability and adaptive capacity of hybrid ecosystems. The following sections explore key aspects of hybrid ecosystem structure and function, first by investigating the role of self-organisation in hybrid ecosystem processes, and then by examining ecological integrity, ecosystem services, and specific self-organizing processes within the anthromes. We conclude by proposing a conceptual framework for navigating self-organisation and ecological integrity in the Anthropocene. This framework provides both theoretical and practical foundations for evaluating ecosystem health, guiding sustainable management strategies, and balancing ecological integrity with human well-being amidst rapid environmental change. The Role of Self-Organisation in Hybrid Ecosystem Processes Self-organisation and imposed organisation are two distinct but interrelated processes that shape the structure, function, and integrity of hybrid ecosystems. Understanding the specific role of self-organisation in ecosystems influenced by external interventions is crucial for assessing the effects of both ecological and anthropogenic factors on life-supporting ecosystems. Self-organisation refers to a spontaneous, bottom-up process where system components interact independently, creating patterns and structures without external control. This phenomenon is observable in natural ecosystems, such as the organisation of ant colonies (Gordon, 1999) or neural activity in the brain (Kelso, 1995). Self-organisation exemplifies the inherent adaptability and resilience of ecosystems as they evolve in response to both internal and external changes (Levin, 1998). In contrast, imposed organisation is a top-down process, in which an external authority designs and controls a system to achieve specific objectives. This approach is typical in human systems, such as agricultural fields, urban ecosystems, planted woodlands, or managed nature reserves, where deliberate interventions are applied to achieve desired outcomes (Cumming, 2011). In the Anthropocene, most ecosystems are hybrids of these two organisational modes. Each ecosystem contains elements that are self-organized and others that are shaped by anthropogenic forces. In healthy, natural systems, self-organized dynamics dominate, while in artificial, anthropogenic systems, imposed organisation prevails. Each system can be characterized based on the proportion of self-organized and imposed elements. For example, a planned orchard reflects imposed organisation with its neatly aligned rows, while the spontaneous spacing of shrubs in a natural landscape, driven by competition for water and nutrients, illustrates self-organisation. Interestingly, both processes can generate similar spatial patterns, yet they arise from fundamentally different mechanisms (Rietkerk et al., 2004). In both natural and hybrid ecosystems, self-organisation manifests through six key ecological processes: (1) Pattern Formation , which enhances spatial resource capture and resilience; (2) Population Self-Regulation , which stabilizes species dynamics; (3) Ecological Feedback Loops , which reinforce stability or drive adaptation; (4) Ecosystem Engineering , which creates critical microhabitats and resource hotspots; (5) Functional Food Web Organisation , which maintains the flow of energy and nutrients; and (6) Functional Redundancy and Response Diversity, which buffer ecosystem functions under disturbances. Together, these processes underpin the functioning and resilience of hybrid ecosystems, integrating self-organized ecological processes with human management. This integration strengthens ecological integrity and ensures the continued delivery of ecosystem services in the Anthropocene. Spatial pattern formation is one of the most observable manifestations of self-organisation in ecological systems. Recurrent spatial arrangements such as shrub patches, vegetation bands, or ”fairy circles” commonly emerge in resource-limited landscapes due to feedback between vegetation growth, water redistribution, and soil characteristics. These patterns, which optimize resource capture and reduce ecological stress, are particularly prominent in arid and semi-arid environments (Rietkerk et al., 2004). The second fundamental self-organisation process is population self-regulation , whereby species regulate their own abundance through feedback mechanisms such as resource limitation, predation, and disease dynamics. These processes prevent uncontrolled population booms or crashes, helping stabilize community composition and its ecosystem functions (Murdoch, 1994). The concept of carrying capacity (or self-thinning) represents a self-organizing mechanism that stabilizes biomass or population levels. Ecological feedback loops , both positive and negative, further regulate system behaviour. Positive feedback reinforces desirable processes, such as vegetation-driven increase in soil moisture, while negative feedbacks, such as predator control over herbivore populations, counteract excesses and promote system stability. This self-organized feedback is essential for understanding ecological thresholds, phase transitions, and regime shifts (Walker et al., 2004). Ecosystem engineering is another key component of self-organisation. It involves organisms that modify the physical environment in ways that affect not only their own survival but also the survival and function of the broader ecosystem. Examples include beavers constructing dams, corals building reef structures, and cyanobacteria forming biological soil crusts that regulate water distribution and soil erosion in drylands (Jones et al., 1994). Emerging self-organizing functions of food webs also play a critical role, as the structure and strength of trophic interactions affect energy flow, nutrient cycling, and system stability. Well-structured food webs exhibit resilience through redundancy and buffering among trophic levels, preventing the collapse of key ecosystem processes, even under perturbation (Thebault & Fontaine, 2010). Lastly, functional redundancy and response diversity describe how multiple species fulfilling similar roles but differing in their sensitivity to environmental change are fundamental to self-organized resilience. Trait diversity ensures that essential ecosystem functions, such as pollination, decomposition, and primary productivity, are maintained even as species decline (Elmqvist et al., 2003; Mori et al., 2013). Response diversity plays a central role in maintaining functional stability, despite natural stressors and anthropogenic impacts. In systems such as agroecosystems, managed rangelands, and semi-natural forests, species that share similar ecological functions but vary in their tolerance to disturbance help maintain the continuity of key services during times of stress (Elmqvist et al., 2003; Oliver et al., 2015). Response diversity enables ecosystems to bridge natural functioning with human needs, ensuring the sustained delivery of services such as pest control, water regulation, and soil stabilization even when certain components of the system are lost. Moreover, response diversity enhances the system’s capacity for self-repair. After disturbances such as drought, fire, or overgrazing, species with differing responses can re-establish functional structures and processes, through self-organisation, without extensive human intervention (Mori et al., 2013). Natural processes within ecosystems reduce the dependence on external input and promote more sustainable forms of land use. In this context, self-organisation acts not only as an ecological principle but also as a practical mechanism for adaptive management, buffering hybrid ecosystems against instability and fostering long-term sustainability. Ultimately, self-organisation, reinforced by response diversity, serves as a foundational mechanism through which hybrid ecosystems retain integrity and functionality in the Anthropocene. Acknowledging and enhancing these self-organizing processes in ecological management can inform more resilient and adaptive strategies for sustaining ecosystem services in an era of unprecedented environmental change. Self-Organizing Processes, Ecological Integrity, and Ecosystem Services of Specific Hybrid Ecosystems: the Anthromes The Anthropocene has brought about the global dominance of hybrid ecosystems that combine natural self-organisation with varying degrees of human management. The anthrome framework, developed by Ellis (Ellis & Ramankutty, 2008), classifies these human-influenced systems based on population density, land use, and ecological structure. Building on this framework, it is crucial to understand how self-organisation sustains ecological integrity and ecosystem services across different anthrome categories, each representing a continuum between natural processes and human intervention. Although human activities dominate croplands , self-organizing processes remain vital. Spatial pattern formation appears through the diversification of cropping systems, such as intercropping and agroforestry, which optimize resource use and stabilize yields (Kremen & Miles, 2012). Population self-regulation occurs through predator-prey dynamics between natural enemies and crop pests (Letourneau et al., 2009). Feedback loops, such as soil health improvement through organic matter recycling, enhance fertility and resilience (Gliessman, 2015). Plant root systems that enhance soil structure and microbial communities act as ecosystem engineers (Lavelle et al., 1997). Food web organisation, by pollinators and decomposers among others, ensures sustained production (Tscharntke et al., 2005). Functional redundancy and response diversity are maintained by cultivating multiple crop varieties with varying tolerances to drought or pests, buffering agricultural systems against climatic variability (Altieri, 1999). Agricultural practices become sustainable when self-organized processes occur, and farmer intervention is minimized. For example, nitrogen supply from a self-organized food web is considered healthier than a continuous supply of artificial nitrogen. The balance between bacteria and fungi, and the control of harmful pests by self-organized food webs, reduce the need for pesticides and antibiotics. Rather than categorizing agriculture as either intensive or organic, we can view agricultural practices along a gradient of self-organisation levels. Thus, self-organisation can be used as a measure of system health, and the level of EI of the agricultural system depends on the extent of self-organisation present. In rangelands , self-organizing vegetation patterns emerge from grazing-mediated feedback and differential resource availability (Rietkerk et al., 2004; Briske et al., 2023). Population self-regulation is observed through herbivore-vegetation interactions that prevent overgrazing under moderate stocking rates (Fuhlendorf et al., 2012). Feedback loops, such as positive feedback between plant cover and soil moisture retention, sustain grassland productivity (Bestelmeyer et al., 2006). Ecosystem engineering by species like prairie dogs and termites alters soil structure, influencing hydrology and plant diversity (Davidson et al., 2012). Functional food webs involving grazers, predators, and scavengers maintain ecosystem energy flow (Polis et al., 1996). Functional redundancy among different grass and shrub species enhances rangeland resilience to drought and grazing pressure (Sasaki et al., 2009). In village landscapes and rural mosaics, self-organisation is evident in patchy mosaics of gardens, fallow fields, and forests that emerge from decentralized land-use decisions (Altieri & Toledo, 2011). Population self-regulation is maintained through local seed networks that promote diverse crop varieties adapted to different microclimates (Zimmerer, 1996). Feedback loops exist in nutrient cycling through composting and livestock integration into farming systems (Tittonell, 2014). Ecosystem engineering is evident in traditional water harvesting infrastructures, which create microhabitats and sustain soil moisture (Reij et al., 1996). Functional food webs, involving native pollinators and pest control organisms, emerge naturally in diversified fields (Perfecto & Vandermeer, 2010). Functional redundancy is preserved through polycultures and traditional crop rotations, enhancing system adaptability (Altieri, 1999). In residential and wildland mosaics , self-organisation is evident in spontaneous succession processes in vacant lots, parks, and urban peripheries (Aronson et al., 2017; Kowarik, 2011). Population self-regulation operates through urban wildlife dynamics, such as predator-prey interactions among small mammals, birds, and their predators (Shochat et al., 2006). Feedback loops regulate green space resilience; for example, vegetation improves soil permeability and mitigates urban runoff (Bolund & Hunhammar, 1999). Tree roots act as ecosystem engineers by stabilizing the soil and providing shade, thereby influencing microclimates (McKinney, 2006). Functional food webs develop in urban settings, involving insects, birds, and mammals that facilitate seed dispersal and pest control (Alberti, 2005). Functional redundancy in urban flora, with diverse plant species adapted to different disturbance regimes, ensures continued green infrastructure services (Kattwinkel et al., 2011). In forest -populated mosaics, spatial patterns form through natural gap dynamics and regeneration after selective logging or small-scale agricultural endeavours (Chazdon, 2003; Chazdon & Guariguata, 2016). Population self-regulation is mediated by density-dependent seedling survival and herbivory (Connell, 1978). Feedback loops involving forest canopy closure and microclimate stabilization help maintain successional trajectories (Pickett & White, 1985). Ecosystem engineering by large trees and animals (e.g., seed dispersal by primates or birds) shapes forest structure and biodiversity (Asner et al., 2010). Functional food webs involving multi-trophic interactions between plants, herbivores, predators, and decomposers maintain nutrient cycling and productivity (Terborgh et al., 2001). Functional redundancy is preserved through species-rich assemblages that ensure functional continuity despite localized disturbances (Mori et al., 2013). Seminatural systems, such as rewilded pastures and abandoned fields undergoing secondary succession, are characterized by natural spatial pattern formation, often influenced by disturbance regimes like fire and grazing (Benayas et al., 2009). Population self-regulation occurs through herbivore browsing pressure, that modulates vegetation dynamics (Bond, 2019). Feedback loops include vegetation-fire interactions that stabilize or shift community composition (Pausas & Keeley, 2009). Large herbivores, such as bison or horses, are an example of ecosystem engineers that modify grassland structure (Sandom et al., 2014). Functional food webs develop through recolonization by trophic guilds, from primary consumers to apex predators (Svenning et al., 2016). Functional redundancy and response diversity are essential in these systems, increasing resilience to climatic variability (Perring et al., 2015). Within all anthrome categories, these six self-organizing processes form the ecological foundation that underpins resilience, ecological integrity, and the provision of ecosystem services. Recognizing, protecting, and enhancing these self-organizing dynamics, particularly through the promotion of functional redundancy and response diversity, offers a pathway toward adaptive and sustainable management of hybrid landscapes in an era of rapid global change. The Role of Self-Organisation in Hybrid Ecosystems’ Function, Sustainability, and Integrity in the Anthropocene In the Anthropocene, hybrid ecosystems, combining elements of self-organisation with human-imposed structures, are the dominant form of terrestrial ecosystems (Ellis and Ramankutty 2008; Kennedy et al., 2019). Understanding the role of self-organisation in these systems is essential for evaluating their ecological integrity, sustainability, and their capacity to maintain critical ecosystem functions under escalating environmental pressures. Despite the impact anthropogenic pressures, hybrid ecosystems continue to rely heavily on self-organizing dynamics for their resilience and functionality. For example, in croplands, soil microbial cycling and plant-pollinator networks persist even under intensive agricultural practices, supporting nutrient cycling, pest control, and soil health (Kremen & Miles, 2012; Gliessman, 2015). In rangelands, spontaneous vegetation pattern formation and grazing feedbacks help maintain forage production and erosion control alongside human management (Briske et al., 2023; Sanderson et al., 2020). In village mosaics, agroecological succession and seed exchange networks foster biodiversity, genetic resilience, and ecosystem service provision (Altieri & Toledo, 2011; Tittonell, 2014). In addition to supporting the internal stability of hybrid systems, self-organisation increases their capacity to deliver critical ecosystem services, including provisioning services (e.g., food, fibre, and fuel), regulating services (e.g., climate moderation, erosion control), supporting services (e.g., nutrient cycling, habitat provision), and cultural services (e.g., traditional knowledge and identity). Moreover, self-organizing processes reduce the dependency of hybrid ecosystems on constant human inputs. Systems that maintain mechanisms such as self-repair through successional dynamics, feedback stabilization, or trophic redundancy demonstrate greater adaptive capacity and long-term sustainability (Walker et al., 2004; Chazdon & Guariguata, 2016). However, challenges remain. Hybrid systems are often vulnerable to management practices that suppress self-organisation such as monoculture expansion, urban sprawl, overgrazing, or fire suppression which can lead to reduced resilience, ecological degradation, and the loss of essential services (Benayas et al., 2009; Kowarik, 2011). Recognizing the intertwined roles of self-organisation and human management offers a practical pathway for adaptive ecosystem management in the Anthropocene. Strategies should not only aim to mitigate human impacts but to enhance and protect self-organizing processes such as fostering landscape heterogeneity, conserving functional redundancy, and promoting multi-scale feedbacks (Sayer et al., 2013; Aronson et al., 2017). In conclusion, self-organisation remains a cornerstone of ecological integrity in hybrid ecosystems. These intrinsic processes must be protected and strengthened, to maintain ecosystem services, promote resilience, and ensure the sustainability of human-environment systems in an era of accelerating global change. Using Self-Organisation as a Quantitative Measure of Hybrid Ecosystem Integrity EI is commonly viewed as a measure of ecosystem health (Leopold, 1949; Karr, 1991). However, its definition, quantification, and application have remained unclear. Historically, the concept of EI focused on preserving pristine environments, primarily within protected areas, to maintain what were considered to be untouched, natural conditions (Hermoso & Clavero, 2013; Andreasen et al., 2001). Aldo Leopold’s seminal work (1949) emphasized the importance of maintaining the integrity of biotic communities to ensure ecosystem stability (Wurzebach & Schultz, 2016), which later became foundational for environmental policies such as the Clean Water Act (Karr, 1991; Cafaro & Primack, 2014). The development of the concept of EI filled an important gap by describing ecosystems’ capacity to maintain valuable services under both internal and external drivers (Bridgewater et al., 2014). EI has been defined as the ability of ecosystems to support the provision of services in the absence of anthropogenic disturbances (Del Leo & Levin, 1997; Muller et al., 2000), or as a “precaution against unspecific ecological risks within the framework of sustainable development” (Karr & Dudley, 1981). EI later replaced the term “supporting ecosystem services” to describe conditions capable of sustaining ecosystem services (Burkhard et al., 2010; Haines-Young & Potschin, 2010; Kandziora et al., 2013); its interpretation was dependent on societal preferences (Kay & Schneider, 1994). When EI is defined solely as the lack of human interference (Hermoso & Clavero, 2013), assessing the actual impact of anthropogenic disturbances on an ecosystem becomes problematic. Relating high EI to a pristine ecosystem creates a circular argument . If an ecosystem begins to include anthropogenic activities, assessing their impact using EI—measured by the presence or absence of anthropogenic activity—is unfeasible. A clearer distinction is needed between the presence of anthropogenic activity and its impact, which is especially relevant when discussing hybrid ecosystems. A hybrid ecosystem can be strongly affected by anthropogenic disturbances while most of its processes and structure are still shaped by self-organisation, or it may be very heavily affected by external forces. The former will be considered to have high EI while the latter has low EI. In this context, we propose an approach that does not focus solely on human interference or needs. Instead, it examines the ecosystem’s intrinsic characteristics, particularly its ability to maintain self-organisation . This approach suggests that when an ecosystem can autonomously organize itself, its integrity remains high, even if it is influenced by anthropogenic factors. Thus, the definition of EI, as proposed by Muller (2005), is framed as ”the ability of a system to maintain self-organisation.” This definition can be applied to hybrid ecosystems by recognizing that these systems, despite human modification, retain self-organizing processes that are integral to their functioning. The key advantage of this approach is that it defines EI based on the system’s ability to maintain organized, functional processes, irrespective of human impacts. This allows for the assessment of anthropogenic impacts by evaluating the degree to which self-organizing processes persist. Therefore, we can grade EI based on the extent of self-organisation present in a system, even if anthropogenic impacts are evident. Self-organisation in hybrid ecosystems can be quantified. Using Muller’s definition of EI (2005), we can quantify the level of self-organisation in a hybrid system and, by extension, determine its EI. A system that is not pristine but continues to exhibit high levels of self-organisation can still be considered to have high integrity, even with anthropogenic influence. Integrity is determined by the ecosystem’s functioning —its ability to maintain core processes such as nutrient cycling, energy flow, and species interactions—rather than by the mere presence of external, human influences. In conclusion, the definition of EI as ”the ability of a system to maintain self-organisation” provides a useful framework for evaluating the integrity of hybrid ecosystems. It shifts the focus from a static, anthropocentric view of EI to one that recognizes the dynamic, self-organizing nature of ecosystems, even under human influence. This approach allows for more nuanced assessments of ecosystem health, particularly in the Anthropocene, where hybrid ecosystems dominate the landscape. Framework for Navigating Self-Organisation and Ecological Integrity of Hybrid Ecosystems in the Anthropocene Navigating the ecological integrity of hybrid ecosystems in the Anthropocene requires a conceptual framework that integrates the role of self-organisation in ecosystem function across different ecosystem types. We propose a roadmap (Fig. 1) that identifies key self-organizing processes, such as spatial pattern formation, population self-regulation, ecological feedback loops, ecosystem engineering, functional food web organisation, and functional redundancy. These self-organizing processes regulate core ecosystem functions, including resource capture, energy flow, nutrient cycling, disturbance recovery, and soil health. These processes are the foundation of whole ecosystem and landscape properties, which in turn ultimately determine the ecological integrity of hybrid ecosystems. The ecological integrity of hybrid ecosystems refers to their ability to maintain essential structure, function, and resilience over time, despite the influences of natural self-organisation and anthropogenic pressures. It reflects the capacity of these systems to sustain ecosystem processes and the services they provide, even under human-imposed organisation. This integrity is determined by the continued presence of self-organizing processes that interact with and complement human intervention, enabling the system to adapt and persist under changing environmental and socio-economic conditions. Building on this roadmap, the framework proposes a targeted evaluation of ecological integrity based on four diagnostic dimensions: (1) the presence and redundancy of key ecosystem functions; (2) the system’s resilience to disturbances such as drought, fire, or land-use change; (3) the spatial structure and heterogeneity of vegetation or habitat; and (4) the level of functional and response diversity among biological communities. These criteria offer a multidimensional lens through which the EI of hybrid systems can be assessed and compared. Importantly, this process-based approach emphasizes the mechanisms by which self-organisation in hybrid ecosystems maintains their capacity to adapt and persist. This aligns with contemporary paradigms in resilience and sustainability science, which prioritize adaptive capacity, feedback regulation, and functional integrity over time. By explicitly incorporating self-organisation into assessments of ecological integrity, this framework enables managers and policymakers to identify which self-organisation processes and ecosystem functions are most critical to conservation or restoration within specific hybrid ecosystems. In doing so, it promotes strategies that work with, rather than override, the inherent ecological processes of ecosystems. Such strategies are particularly vital in the Anthropocene, where most terrestrial ecosystems are hybrid and face mounting socio-ecological challenges. The next stage is to quantify self-organisation in an ecosystem or to quantify the relative impact of each force in an ecosystem driven by self-organisation versus imposed organisation. Using the developed quantitative tools, one can assess the integrity of an ecosystem and thus improve management practices. For example, various conservation tools can be used on hybrid systems to assess the efficiency of each tool. Conclusions The profound human-driven transformations characteristic of the Anthropocene have led to the world-wide dominance of hybrid ecosystems, in which natural self-organisation processes interact with human-imposed systems. These hybrid ecosystems play a critical role in sustaining ecosystem functions and human well-being. This paper has demonstrated that self-organisation is a fundamental driver of EI in hybrid ecosystems by identifying and analysing core self-organizing processes—specifically, spatial pattern formation, population self-regulation, ecological feedback loops, ecosystem engineering, functional food web organisation, and functional redundancy. This paper shows how these processes underpin essential ecosystem functions across various hybrid ecosystem (anthrome) categories. We emphasize that understanding the degree and nature of self-organisation within hybrid ecosystems is essential for assessing their ecological integrity and sustainability. The proposed conceptual framework for evaluating ecological integrity integrates these self-organizing processes, mapping their contributions to hybrid ecosystem structure and function. We argue that EI in these ecosystems should not be defined solely by the extent of human impact, but by the continued presence and interaction of self-organizing processes with human interventions. Moreover, the framework provides a multidimensional approach to assessing EI, emphasizing diagnostic criteria such as the redundancy of key ecosystem functions, resilience to disturbances, spatial structure, and functional and response diversity among biological communities. These dimensions allow for context-sensitive assessments of hybrid ecosystems and offer a dynamic, process-based approach for monitoring their integrity. Using self-organisation can be a useful tool to asses EI in all hybrid ecosystems, and can be an especially useful tool if we quantify the self-organisation processes in hybrid ecosystems. The importance of this framework lies in its ability to guide adaptive management strategies that enhance the inherent ecological processes of hybrid ecosystems. In an era of rapid environmental change, fostering resilience through self-organisation is essential for ensuring the long-term sustainability of ecosystems and the continued provision of ecosystem services. In conclusion, the role of self-organisation in hybrid ecosystems’ ecological integrity is crucial, and acknowledging and reinforcing these processes is vital for promoting sustainability in the Anthropocene. ACKNOWLEDGMENTS We thank the Israel Science Foundation (ISF) for funding this research (grant number 964/14) and the Ministry of Science. We also thank Michelle Finzi for her editing assistance. Data Accessibility Statement Data sharing not applicable to this article as no datasets were generated or analysed during the current study viii. Figure legends and embedded figures Figure 1 : Framework for Navigating Self-Organisation and Ecological Integrity of Hybrid Ecosystems in the Anthropocene . This flowchart presents a comprehensive, process-oriented approach to assessing and managing the EI of hybrid ecosystems. The framework begins by identifying key self-organizing processes and linking them to core ecosystem functions that regulate ecosystem and landscape structure. The resulting ecosystem properties ultimately determine the ecological integrity of hybrid ecosystems. Key diagnostic dimensions for assessing ecological integrity are also proposed. These navigating processes are mapped to specific anthrome categories, reflecting the diversity of human-imposed organisation within hybrid ecosystems. The framework emphasizes the critical role of self-organisation in maintaining the resilience and functionality of hybrid ecosystems. Information & Authors Information Version history V1 Version 1 13 August 2025 Peer review timeline Published Ecology and Evolution Version of Record 8 Apr 2026 Published Copyright This work is licensed under a Non Exclusive No Reuse License. Collection Ecology and Evolution Keywords ecosystem ecosystem ecology ecosystem function terrestrial theoretical theory Authors Affiliations Elli Groner 0000-0002-7843-3047 [email protected] Dead Sea and Arava Science Center View all articles by this author Aviva Peeters Negev School of Architecture, SCE Shamoon College of Engineering View all articles by this author Moshe Shachak Ben-Gurion University of the Negev View all articles by this author Metrics & Citations Metrics Article Usage 356 views 211 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Elli Groner, Aviva Peeters, Moshe Shachak. Ecological Integrity of Hybrid Ecosystems in the Anthropocene: The Impact of Self-Organisation on Function and Sustainability. Authorea . 13 August 2025. DOI: https://doi.org/10.22541/au.175508139.98145484/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. 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