Introduction
Mediterranean climate regions (hereafter ‘Med-regions’), with unusual climatic conditions of mild wet winters and warm to hot dry summers (Lionello et al. Reference Lionello, Malanotte-Rizzoli and Boscolo2006), are biodiversity hotspots that have historically been exposed to anthropogenic impacts (Fig. 1; Myers et al. Reference Myers, Mittermeier, Mittermeier, da Fonseca and Kent2000, Cramer et al. Reference Cramer, Guiot, Fader, Garrabou, Gattuso and Iglesias2018) and thus are ideal open laboratories for global change research (Underwood et al. Reference Underwood, Viers, Klausmeyer, Cox and Shaw2009, Lejeusne et al. Reference Lejeusne, Chevaldonné, Pergent-Martini, Boudouresque and Pérez2010). Research on Med-regions has been conducted at various biological scales, ranging from the genetic (Aurelle et al. Reference Aurelle, Thomas, Albert, Bally, Bondeau and Boudouresque2022, Fady et al. Reference Fady, Esposito, Abulaila, Aleksic, Alia and Alizoti2022) and species level (Heywood Reference Heywood2014, Bianchi et al. Reference Bianchi, Caroli, Guidetti and Morri2018, Zenetos & Galanidi Reference Zenetos and Galanidi2020) to ecological communities (Angulo et al. Reference Angulo, Boulay, Ruano, Tinaut and Cerdá2016, Panitsa et al. Reference Panitsa, Iliopoulou and Petrakis2021), and across freshwater (Gasith & Resh Reference Gasith and Resh1999, Bonada & Resh Reference Bonada and Resh2013), marine (Coll et al. Reference Coll, Piroddi, Steenbeek, Kaschner, Lasram and Aguzzi2010, Gianni et al. Reference Gianni, Bartolini, Airoldi, Ballesteros, Francour and Meinesz2013, Darmaraki et al. Reference Darmaraki, Somot, Sevault, Nabat, Narvaez and Cavicchia2019) and terrestrial (Lavorel et al. Reference Lavorel, Canadell, Rambal and Terradas1998, Underwood et al. Reference Underwood, Viers, Klausmeyer, Cox and Shaw2009, Doblas-Miranda et al. Reference Doblas-Miranda, Alonso, Arnan, Bermejo, Brotons and de las Heras2017) ecosystems. Conservation efforts have been largely dedicated to reducing threats, including overfishing and habitat fragmentation, or predicting the effects of changing fire and heatwave regimes. Conversely, few integrative studies from different areas of knowledge exist, including comparisons amongst global Med-regions (e.g., Mediterranean and California plant communities; Quezel & Berbéro Reference Quezel and Barbéro1989, Medail & Quezel Reference Medail and Quezel1997, Marr et al. Reference Marr, Marchetti, Olden, García-Berthou, Morgan and Arismendi2010) or across ecosystems (e.g., marine and terrestrial biodiversity in the Mediterranean Basin; Aurelle et al. Reference Aurelle, Thomas, Albert, Bally, Bondeau and Boudouresque2022).
Terrestrial, freshwater and marine ecosystems in Med-regions are subject to natural and extreme climate disturbances, including wildfires, floods, droughts and heatwaves (Castri & Mooney Reference Castri and Mooney1973). To cope with these disturbances, species have evolved with similar life-history strategies regardless of Med-region, including, for instance, sclerophyllous leaves or animal aestivation to avoid the effects of the droughts (Fig. 2; David et al. Reference David, Henriques, Kurz-Besson, Nunes, Valente and Vaz2007, Quero et al. Reference Quero, Sterck, Martínez-Vilalta and Villar2011, Bussotti et al. Reference Bussotti, Ferrini, Pollastrini and Fini2014). Given these adaptations to natural disturbances, Med-region ecosystems could also be more tolerant of human disturbances, have wider safe operating spaces (i.e., situations where ecosystems can support essential resources and functions for human needs and well-being, as defined by Rockström et al. Reference Rockström, Steffen, Noone, Persson, Chapin and Lambin2009) and show greater resilience than non-Mediterranean ecosystems (Fox & Fox Reference Fox, Fox, Dell, Hopkins and Lamont1986, Bonada et al. Reference Bonada, Dolédec and Statzner2007, Helman et al. Reference Helman, Lensky, Yakir and Osem2017). These characteristics should allow Med-region ecosystems to react rapidly to overcome ecosystem degradation (Fig. 2b; Lavorel et al. Reference Lavorel, Canadell, Rambal and Terradas1998, Underwood et al. Reference Underwood, Viers, Klausmeyer, Cox and Shaw2009).
Forecasting the dynamics of ecosystems’ responses to changing environmental conditions is challenging due to the difficulty in predicting tipping points (i.e., points at which ecosystems transition from one stable state to another), especially given the uncertainty of ecological baseline conditions (Moore Reference Moore2018). For instance, in the Mediterranean Basin, megafires might shift terrestrial ecosystems towards vegetation formations characteristic of more arid regions (Piñol et al. Reference Piñol, Castellnou and Beven2007, San-Miguel-Ayanz et al. Reference San-Miguel-Ayanz, Moreno and Camia2013). Similarly, high grazing pressure in other terrestrial Med-region ecosystems pushes biotic integrity towards the edge of extinction with decreased canopy cover and high erosion (Kéfi et al. Reference Kéfi, Rietkerk, Alados, Pueyo, Papanastasis, ElAich and de Ruiter2007). Increased aridity can then lead to irreversible desertification, in which the possibility of recovery is likely to be low. In freshwater ecosystems, reduced amounts of precipitation and falling groundwater levels (e.g., 30% rainfall reduction in spring/summer by 2080; MedECC Reference Cramer, Guiot and Marini2020) can reduce the longevity of the hydroperiods of temporary ponds and intermittent streams (Zacharias & Zamparas Reference Zacharias and Zamparas2010). In marine ecosystems, the depletion of sea urchin predators caused by chronic overfishing has been one of the main drivers of macroalgal collapse in the Mediterranean Sea (Ling et al. Reference Ling, Scheibling, Rassweiler, Johnson, Shears and Connell2015). To predict future conditions, it is essential to establish clear ecological baselines and recognize significant changes in ecosystem states.
In this work, we define integrative research as the interdisciplinary approach from multiple fields of study with common concepts and metrics (Winder Reference Winder, Tress, Tress, Van der Valk and Fry2003, Tress et al. Reference Tress, Tress and Fry2005) that enables a better understanding of the general mechanisms underlying ecosystems. In Med-regions, integrative research could provide more effective conservation strategies with a suite of resilience-based management approaches, including developing incentives that encourage sustainability, identifying ecological baselines and protecting key species and ecosystems (Abiven et al. Reference Abiven, Altermatt, Backhaus, Deplazes-Zemp, Furrer and Korf2017). Previous integrative studies in Med-regions have elucidated common ecological patterns, contributing to the enhancement of conservation practices. A synthesis by Bonada & Resh (Reference Bonada and Resh2013) based on 22 papers addressed diversity patterns and conservation practices but only regarding Mediterranean rivers and streams. Lavorel et al. (Reference Lavorel, Canadell, Rambal and Terradas1998) provided the first roadmap for conducting global change research in Mediterranean forests and pointed out the need for ‘appropriate coordination and synthesis of that research’. More recently, a group of scientists with different scientific backgrounds has coordinated an exhaustive report emphasizing the potential hazards of climate and environmental changes in the Mediterranean Basin (Cramer et al. Reference Cramer, Guiot, Fader, Garrabou, Gattuso and Iglesias2018, Hassoun et al. Reference Hassoun, Guiot, Marini and Cramer2021). In the current work, building on previous efforts highlighting the need to promote integrative research in Med-regions, we advocate for the exploration of cross-ecosystem generalities and common resilience mechanisms (e.g., tipping points) in Med-regions.
Here, we aim to promote integrative research across ecosystems in Med-regions, which are heavily threatened by the most significant factors raised in the World Scientists’ Warning to Humanity manifesto (e.g., increased temperature, human population density; Ripple et al. Reference Ripple, Wolf, Newsome, Galetti, Alamgir and Crist2017, Reference Ripple, Wolf, Gregg, Rockström, Newsome and Law2023). Although integrative research covering Med-regions globally was scarce at the time of the first World Scientists’ Warning, more comprehensive ecological knowledge and datasets are now available, allowing for the identification of ecological commonalities across Med-regions (e.g., Aurelle et al. Reference Aurelle, Thomas, Albert, Bally, Bondeau and Boudouresque2022, Albassatneh et al. Reference Albassatneh, Dubuisson, Escudero, Fady, Ponger and Muller2023, Herrera et al. Reference Herrera, Núñez, Valverde and Alonso2023). Thus, we first propose a call to action to the research community for integrative studies in Med-regions. We then offer recommendations for improving such integrative studies, including leveraging biodiversity data, promoting working groups in research syntheses and testing theoretical frameworks.
Integrative studies in Med-regions
We suggest that there are three main types of integrative studies at different spatial scales, from local to global, which would enable and facilitate an integrative research framework in Med-regions. We propose such studies to represent (1) integrative research across ecosystems, (2) integrative research across Med-regions and (3) integrative research across political boundaries.
Integrative research across ecosystems
The connectivity amongst different ecosystem types plays a vital role in shaping nutrient exchanges, community dynamics and food web structure (Polis et al. Reference Polis, Anderson and Holt1997). This is because ecosystems are never isolated in space but interact through the movement of matter, nutrients and organisms across habitat boundaries (Gravel et al. Reference Gravel, Guichard, Loreau and Mouquet2010). Few studies on Med-regions have focused on the connectivity amongst ecosystem types (Bastow et al. Reference Bastow, Sabo, Finlay and Power2002, Rundio & Lindley Reference Rundio and Lindley2012, Orihuela-Torres et al. Reference Orihuela-Torres, Pérez-García, Sánchez-Zapata, Botella and Sebastián-González2022). For instance, Orihuela-Torres et al. (Reference Orihuela-Torres, Pérez-García, Sánchez-Zapata, Botella and Sebastián-González2022) found in Mediterranean ponds (Spain) that the consumption of carp carcasses by terrestrial vertebrate scavengers serves as a link between the aquatic and terrestrial ecosystems. Bastow et al. (Reference Bastow, Sabo, Finlay and Power2002) showed in a Mediterranean river in California that large populations of terrestrial grasshoppers can be sustained due to their high rates of consumption of algae, providing an alternative pathway for energy exchange between rivers and riparian habitats. We argue that a priority in integrative research within Med-regions would be to identify cross-ecosystem boundaries that encompass aquatic–terrestrial linkages (e.g., coastal dune–sea, pond–surrounding landscape, stream–riparian forest) and to integrate knowledge from diverse disciplines, placing a specific focus on establishing ecological baselines (Graham et al. Reference Graham, Wary, Calcagni, Cisneros, de Luca and Gorostiza2023). The prioritization of zones with the highest level of inter-ecosystem connectivity (i.e., aquatic–terrestrial or freshwater–marine linkages) would complement earlier perspectives, in which priority-setting relied more heavily on iconic or single species within an ecosystem (Panou et al. Reference Panou, Jacobs and Panos1993, Alessandro & Antonello Reference Alessandro and Antonello2010, Palomares et al. Reference Palomares, Rodríguez, Revilla, López-Bao and Calzada2011).
Integrative research across Med-regions
Although extensive research has been conducted to understand the impacts of global change factors on biodiversity in the Mediterranean Basin (e.g., Bonada & Resh Reference Bonada and Resh2013, Doblas-Miranda et al. Reference Doblas-Miranda, Alonso, Arnan, Bermejo, Brotons and de las Heras2017), limited knowledge applies to less developed Med-regions such as central Chile (Figueroa et al. Reference Figueroa, Bonada, Guevara, Pedreros, Correa-Araneda, Díaz and Ruiz2013). This leads to a piecemeal understanding of Med-regions and a less effective integration of research from different Med-regions worldwide. An operational framework for addressing this can be accomplished through the identification of other effective area-based conservation measures (i.e., those extending beyond traditional protected areas; Dudley Reference Dudley, Jonas, Nelson, Parrish, Pyhälä, Stolton and Watson2018) and the implementation of successful conservation strategies aimed at establishing ecological baselines (Dudley et al. Reference Dudley, Jonas, Nelson, Parrish, Pyhälä, Stolton and Watson2018, Petza et al. Reference Petza, Chalkias, Koukourouvli, Coll, Vassilopoulou and Karachle2019). Moreover, the implementation of research programmes or new tools from one region may improve the knowledge of the ecological status of ecosystems in other regions. For instance, the European Water Framework Directive, drawing on the river knowledge in the northern Mediterranean Basin, has already been employed to facilitate the implementation of conservation plans for Chilean river ecosystems (Fuster et al. Reference Fuster, Escobar, Lillo and de la Fuente2015). The Nature Futures Framework developed by the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IBPES; available at www.ipbes.net) may improve the effectiveness of conservation plans and actions.
Integrative research across political boundaries
Because biological diversity exists in a combination of public and private lands, the designation of protected areas (e.g., national parks, nature reserves; Dudley Reference Dudley2008) is influenced by diverse and complex socio-economic and political factors. For instance, in Chile, only 10% of the Mediterranean region is formally protected (i.e., as a national asset by the government), and the rest belongs to private lands (Simonetti Reference Simonetti1999, Pauchard & Villarroeal Reference Pauchard and Villaroel2002). Moreover, there are some Med-regions with a high percentage of protected land (e.g., over 39% in southern and south-western Australia; Fig. 3) that can still fail to coordinate cross-boundary conservation because this often requires extra logistical support and resources compared to local initiatives (Kark et al. Reference Kark, Levin, Grantham and Possingham2009). Quantitative conservation geography is a subfield in biogeography that integrates elements of human geography, such as cultures and people, and the environmental context (Minin et al. Reference Minin, Correia and Toivonen2022). Quantitative conservation geography aims to implement conservation actions in order to mitigate threats and support sustainable people–nature interactions (Minin et al. Reference Minin, Correia and Toivonen2022), and it can ensure the designation of more integrative research by identifying priority areas with high levels of endemism (Kier et al. Reference Kier, Kreft, Lee, Jetz, Ibisch and Nowicki2009), establish biological corridors (Gonçalves et al. Reference Gonçalves, Martínez-Freiría, Crochet, Geniez, Carranza and Brito2018) and foster collaboration between public services and private landowners (Cortés-Capano et al. Reference Cortés-Capano, Toivonen, Soutullo, Fernández, Dimitriadis, Garibotto-Carton and Di Minin2020). We suggest that moving towards quantitative conservation geography, with a strong emphasis on data acquisition from multiple sources such as remote sensing or field surveys rather than being constrained by political boundaries (Médail & Diadema Reference Médail and Diadema2009, Cheikh Albassatneh et al. Reference Cheikh Albassatneh, Escudero, Monnet, Arroyo, Bacchetta and Bagnoli2021), will enhance the process of identifying ecological baselines and the formulation of effective conservation strategies. Similarly, international organizations, including non-governmental or multilateral development entities, can play a crucial role in incorporating global policies in Med-regions by facilitating effective transboundary coordination and flexible decision-making within global protected area planning frameworks. For instance, the International Organization for Standardization (2018) emphasizes the importance of developing standardized research designs and methodologies to establish baseline references for ecosystem states.
Identifying ecological baselines
Developing integrative studies would improve our comprehension of ecological baselines in Med-regions holistically. Ecological baselines are essential for reliably measuring biodiversity change and assisting management goals (Mihoub et al. Reference Mihoub, Henle, Titeux, Brotons, Brummitt and Schmeller2017). Although Med-regions are already heavily altered by global change impacts, long-term biodiversity monitoring can track the trends and magnitudes of future biodiversity changes and disentangle anthropogenic effects from the impacts of natural factors. Additionally, prior to the original work on biodiversity monitoring, information regarding anthropogenic pressures might not have been accessible, affecting views of the ecological baselines (Pauly Reference Pauly1995). Promoting integrative research across Med-regions will help increase the chances of locating the most pristine areas and using them as ecological baselines for those that have experienced long-term anthropogenic disturbances.
Implications and future directions
Our purpose is to engage the research community in the implementation of integrative research by promoting potential studies that focus on aquatic–terrestrial linkages across different Med-regions and across political boundaries. Despite the strong ecological similarities across Med-regions, to date there has been little integrative research amongst them (but see Lavorel et al. Reference Lavorel, Canadell, Rambal and Terradas1998, Bonada & Resh 2003). We point to several major impediments that can explain why inconsistencies still exist in management strategies across Med-regions worldwide, including lags in data exchange, barriers to an effective working synthesis and a lack of initiatives from international organizations. Moreover, such impediments can also result in unresolved fundamental questions regarding how Mediterranean biodiversity will respond to impacts of global change. Future studies could address these research gaps by investigating where and why biological diversity responds most significantly to global change impacts and identifying the most successful management regulations across Med-regions. Therefore, we emphasize the importance of developing research tools to implement integrative studies. Given the strong ecological similarities amongst Med-regions, a joint research strategy could be beneficial for ensuring the effective conservation and management of Med-regions worldwide experiencing the impacts of global change.
Apart from advocating for an interdisciplinary approach in integrative research, it is also worth considering the cooperation between stakeholders (e.g., managers or policymakers) and researchers, focusing on a transdisciplinary approach. This could allow for the implementation of better decisions, the quicker formulation of problem resolutions and the identification of knowledge gaps in the process of planning integrative studies (Voinov & Bousquet Reference Voinov and Bousquet2010, Cimpoiasu et al. Reference Cimpoiasu, Dowdeswell-Downey, Evans, McCloskey, Rose and Sayer2021). Stakeholders harbour local ecological knowledge that researchers could use to frame their theoretical hypotheses by considering the context dependency of the local environment (Jongman & Padovani Reference Jongman and Padovani2006). Similarly, stakeholders offer a historical perspective on the local environmental changes that researchers can exploit when conducting long-term analyses. Collectively, coordinated research projects involving stakeholders may yield mutual benefits for forecasting the dynamics of ecosystems’ responses to changing environmental conditions and for developing scientific-based practices and policies.
Future studies in Med-regions could be framed within the three types of integrative studies outlined above, which could serve to establish ecological baseline conditions and aid in predicting tipping points through an increased understanding of causal factors (Houlahan et al. Reference Houlahan, McKinney, Anderson and McGill2017). We believe that these recommendations could serve as a starting point for improving our understanding of the responses of Med-region ecosystems to natural and anthropogenic disturbances and for identifying ecological baselines. One promising avenue involves applying and testing the resilience of Med-region ecosystems in conjunction with existing mechanistic theoretical frameworks (e.g., the theory of island biogeography; McArthur & Wilson Reference MacArthur and Wilson1967). This interdisciplinary approach could help to elucidate complex phenomena, including the non-equilibrium aspect of Med-region ecosystems due to natural disturbances, as well as help to identify key drivers that might influence the stability and adaptive capacity of Med-regions. Another important task for future research is leveraging the availability of large spatial and temporal biodiversity datasets in Med-regions. The increasing number of biodiversity datasets offers exciting prospects and challenges for documenting and forecasting changes in the biological diversity and resilience of Med-region ecosystems (Dornelas et al. Reference Dornelas, Antao, Moyes, Bates, Gould and Henshaw2018, Comte et al. Reference Comte, Carvajal-Quintero, Tedesco, Giam, Brose and Erös2021). Synthesis centres offer the opportunity to bring together experts from diverse backgrounds, disciplines and perspectives to collectively address big questions in ecology using these existing biodiversity datasets (Rodrigo et al. Reference Rodrigo, Alberts, Cranston, Kingsolver, Lapp and McClain2013, Baron et al. Reference Baron, Specht, Garnier, Bishop, Campbell and Davis2017). Overall, such approaches would increase the knowledge available for addressing many fundamental and applied questions in Med-regions, facilitating cross-validations and providing a clearer picture of remaining gaps, thereby guiding future sampling and research efforts.
Conclusion
Integrative research is key to detecting similarities that bridge gaps in our understanding of the nature of different ecosystem types in Med-regions. Here, we highlight this in the context of the claim presented in the 1992 World Scientists’ Warning to Humanity manifesto (Ripple et al. Reference Ripple, Wolf, Newsome, Galetti, Alamgir and Crist2017, Reference Ripple, Wolf, Gregg, Rockström, Newsome and Law2023) that there is an urgent need for integrative research to avert the consequences of irreversible ecosystem changes. Considering that most Med-regions feature highly heterogeneous landscapes and seascapes and their species show considerable genetic adaptation to stressors (Rundel et al. Reference Rundel, Arroyo, Cowling, Keeley, Lamont, Pausas and Vargas2018, Aurelle et al. Reference Aurelle, Thomas, Albert, Bally, Bondeau and Boudouresque2022), Med-regions contain unique sets of ecosystems that might exhibit higher resilience capacities to natural disturbance and wider safe operation spaces than more temperate (Bonada et al. Reference Bonada, Dolédec and Statzner2007) or humid (Helman et al. Reference Helman, Lensky, Yakir and Osem2017) ecosystems. However, crossing certain biophysical thresholds could have disastrous consequences for ecosystem integrity, to the point of pushing Med-regions outside their stable zone (e.g., a desertification process). Promoting integrative research could therefore help us to elucidate ecological generalities, promote knowledge–action outcomes and foster multiple disciplinary approaches to anticipate tipping points before the appearance of irreversible ecosystem changes in Med-regions.
Acknowledgements
We are grateful for feedback from Matthew RD Cobain and we thank Nicholas Polunin and three anonymous reviewers for their valuable comments on previous versions of this manuscript.
Financial support
IA is supported by the Programa Talento (Comunidad de Madrid; project FRESCO: 2022-T1/AMB-24100). EB is a Serra Húnter fellow.
Competing interests
The authors declare none.
Ethical standards
None.