Resilience dynamics and productivity-driven shifts in the marine communities of the Western Mediterranean Sea.

Ecological resilience has become a conceptual cornerstone bridging ecological processes to conservation needs. Global change is increasingly associated with local changes in environmental conditions that can cause abrupt ecosystem reorganizations attending to system-specific resilience fluctuations with time (i.e. resilience dynamics). Here we assess resilience dynamics associated with climate-driven ecosystems transitions, expressed as changes in the relevant contribution of species with different life-history strategies, in two benthopelagic systems. We analysed data from 1994 to 2019 coming from a scientific bottom trawl survey in two environmentally contrasting ecosystems in the Western Mediterranean Sea-Northern Spain and Alboran Sea. Benthopelagic species were categorized according to their life-history strategies (opportunistic, periodic and equilibrium), ecosystem functions and habitats. We implemented an Integrated Resilience Assessment (IRA) to elucidate the response mechanism of the studied ecosystems to several candidate environmental stressors and quantify the ecosystems' resilience. We demonstrate that both ecosystems responded discontinuously to changes in chlorophyll-a concentration more than any other stressor. The response in Northern Spain indicated a more overarching regime shift than in the Alboran Sea. Opportunistic fish were unfavoured in both ecosystems in the recent periods, while invertebrate species of short life cycle were generally favoured, particularly benthic species in the Alboran Sea. The study illustrates that the resilience dynamics of the two ecosystems were mostly associated with fluctuating productivity, but subtle and long-term effects from sea warming and fishing reduction were also discernible. Such dynamics are typical of systems with wide environmental gradient such as the Northern Spain, as well as systems with highly hydrodynamic and of biogeographical complexity such as the Alboran Sea. We stress that management should become more adaptive by utilizing the knowledge on the systems' productivity thresholds and underlying shifts to help anticipate both short-term/less predictable events and long-term/expected effects of climate change.

Evidence of climate-driven regime shifts in the Aegean Sea's demersal resources: A study spanning six decades.

Climate change (CC) can alter the configuration of marine ecosystems; however, ecosystem response and resilience to change are usually case-specific. The effect of CC on the demersal resources of the Aegean Sea (east Mediterranean Sea) was investigated during the past six decades applying a combination of multivariate analysis, non-additive modeling and the Integrated Resilience Assessment (IRA) framework. We focused on the study of: (i) the biological "system" complex, using proxies of biomass (landings per unit of capacity) for 12 demersal taxa, and (ii) the environmental "stressor" complex, described by 12 abiotic variables. Pronounced changes have occurred in both the environmental and biological system over the studied period. The majority of the environmental stressors exhibited strikingly increasing trends (temperature, salinity, primary production indices) with values started exceeding the global historical means during late 1980s-early 1990s. It is suggested that the biological system exhibited a discontinuous response to CC, with two apparently climate-induced regime shifts occurring in the past 25 years. There is evidence for two-fold bifurcations and four tipping points in the system, forming a folded stability landscape with three basins of attraction. The shape of the stability landscape for the Aegean Sea's biological system suggests that while the initial state (1966-1991) was rather resilient to CC, absorbing two environmental step-changes, this was not the case for the two subsequent ones (intermediate: 1992-2002; recent: 2003-2016). Given the current trajectory of environmental change, it is highly unlikely that the biological system will ever return to its pre-1990s state, as it is entering areas of unprecedented climatic conditions and there is some evidence that the system may be even shifting toward a new state. Our approach and findings may be relevant to other marine areas of the Mediterranean and beyond, undergoing climate-driven regime shifts, and can assist to their adaptive management.

An Integrated Traits Resilience Assessment of Mediterranean fisheries landings.

An increasing number of studies have been examining the functional configuration of biological communities or ecosystems using biological traits. Here, we investigated the temporal dynamics and resilience of the traits composition in Mediterranean fisheries landings over 31 years (1985-2015). We transcribed the FAO Mediterranean landings dataset for 101 marine species into a dataset of 23 traits related to the life cycle, distribution, ecology and behaviour. Mediterranean mean Sea Surface Temperature (SST) was evaluated as a potential driver of the traits composition. Trait dynamics were evaluated both individually and holistically by developing an Integrated Traits Resilience Assessment (ITRA). ITRA is a variation of the Integrated Resilience Assessment (IRA), a method to infer resilience dynamics and build stability landscapes of complex natural systems. Changes in landings trait dynamics were documented both for individual traits and for the entire traits 'system', and a relevant regime shift was detected in the second half of the 1990s. The traits system switched to higher optimal temperature, more summer spawning, shorter life span, smaller maximum size, shallower optimal depth and planktivorous diet. This shift was found to be a lagged discontinuous response to sea warming, which gradually eroded the resilience of the original state of the traits system, leading it into a new basin of attraction. The inclusion of ecological/response traits (related to environmental preferences) in our analyses indicates potential mechanisms that explain the observed shift, while changes in functional/effect traits indicate potential impacts on ecosystem functioning. Our findings suggest that changes in the Mediterranean ecosystems are evidently larger than previously thought, with profound implications for the management of this highly impacted sea. ​.

Resilience and regime shifts in a marine biodiversity hotspot.

Complex natural systems, spanning from individuals and populations to ecosystems and social-ecological systems, often exhibit abrupt reorganizations in response to changing stressors, known as regime shifts or critical transitions. Theory suggests that such systems feature folded stability landscapes with fluctuating resilience, fold-bifurcations, and alternate basins of attraction. However, the implementation of such features to elucidate response mechanisms in an empirical context is scarce, due to the lack of generic approaches to quantify resilience dynamics in individual natural systems. Here, we introduce an Integrated Resilience Assessment (IRA) framework: a three-step analytical process to assess resilience and construct stability landscapes of empirical systems. The proposed framework involves a multivariate analysis to estimate holistic system indicator variables, non-additive modelling to estimate alternate attractors, and a quantitative resilience assessment to scale stability landscapes. We implement this framework to investigate the temporal development of the Mediterranean marine communities in response to sea warming during 1985-2013, using fisheries landings data. Our analysis revealed a nonlinear tropicalisation of the Mediterranean Sea, expressed as abrupt shifts to regimes dominated by thermophilic species. The approach exemplified here for the Mediterranean Sea, revealing previously unknown resilience dynamics driven by climate forcing, can elucidate resilience and shifts in other complex systems.

Resilience and tipping points of an exploited fish population over six decades.

Complex natural systems with eroded resilience, such as populations, ecosystems and socio-ecological systems, respond to small perturbations with abrupt, discontinuous state shifts, or critical transitions. Theory of critical transitions suggests that such systems exhibit fold bifurcations featuring folded response curves, tipping points and alternate attractors. However, there is little empirical evidence of fold bifurcations occurring in actual complex natural systems impacted by multiple stressors. Moreover, resilience of complex systems to change currently lacks clear operational measures with generic application. Here, we provide empirical evidence for the occurrence of a fold bifurcation in an exploited fish population and introduce a generic measure of ecological resilience based on the observed fold bifurcation attributes. We analyse the multivariate development of Barents Sea cod (Gadus morhua), which is currently the world's largest cod stock, over six decades (1949-2009), and identify a population state shift in 1981. By plotting a multivariate population index against a multivariate stressor index, the shift mechanism was revealed suggesting that the observed population shift was a nonlinear response to the combined effects of overfishing and climate change. Annual resilience values were estimated based on the position of each year in relation to the fitted attractors and assumed tipping points of the fold bifurcation. By interpolating the annual resilience values, a folded stability landscape was fit, which was shaped as predicted by theory. The resilience assessment suggested that the population may be close to another tipping point. This study illustrates how a multivariate analysis, supported by theory of critical transitions and accompanied by a quantitative resilience assessment, can clarify shift mechanisms in data-rich complex natural systems.

The alarming decline of Mediterranean fish stocks.

In recent years, fisheries management has succeeded in stabilizing and even improving the state of many global fisheries resources [1-5]. This is particularly evident in areas where stocks are exploited in compliance with scientific advice and strong institutional structures are in place [1, 5]. In Europe, the well-managed northeast (NE) Atlantic fish stocks have been recovering in response to decreasing fishing pressure over the past decade [3-6], albeit with a long way to go for a universal stock rebuild [3, 7]. Meanwhile, little is known about the temporal development of the European Mediterranean stocks, whose management relies on input controls that are often poorly enforced. Here, we perform a meta-analysis of 42 European Mediterranean stocks of nine species in 1990-2010, showing that exploitation rate has been steadily increasing, selectivity (proportional exploitation of juveniles) has been deteriorating, and stocks have been shrinking. We implement species-specific simulation models to quantify changes in exploitation rate and selectivity that would maximize long-term yields and halt stock depletion. We show that stocks would be more resilient to fishing and produce higher long-term yields if harvested a few years after maturation because current selectivity is far from optimal, especially for demersal stocks. The European Common Fisheries Policy that has assisted in improving the state of NE Atlantic fish stocks in the past 10 years has failed to deliver similar results for Mediterranean stocks managed under the same policy. Limiting juvenile exploitation, advancing management plans, and strengthening compliance, control, and enforcement could promote fisheries sustainability in the Mediterranean.