Resilience assessment in complex natural systems.

Ecological resilience is the capability of an ecosystem to maintain the same structure and function and avoid crossing catastrophic tipping points (i.e. undergoing irreversible regime shifts). While fundamental for management, concrete ways to estimate and interpret resilience in real ecosystems are still lacking. Here, we develop an empirical approach to estimate resilience based on the stochastic cusp model derived from catastrophe theory. The cusp model models tipping points derived from a cusp bifurcation. We extend cusp in order to identify the presence of stable and unstable states in complex natural systems. Our Cusp Resilience Assessment (CUSPRA) has three characteristics: (i) it provides estimates on how likely a system is to cross a tipping point (in the form of a cusp bifurcation) characterized by hysteresis, (ii) it assesses resilience in relation to multiple external drivers and (iii) it produces straightforward results for ecosystem-based management. We validate our approach using simulated data and demonstrate its application using empirical time series of an Atlantic cod population and marine ecosystems in the North Sea and the Mediterranean Sea. We show that Cusp Resilience Assessment is a powerful method to empirically estimate resilience in support of a sustainable management of our constantly adapting ecosystems under global climate change.

Fish distribution and behaviour with regard to the time of day and the anthropogenic structural modification of the shallow littoral.

The shallow littoral zone is under a variety of environmental drivers and anthropogenic pressures. As these factors are highly dynamic, they may affect the distribution and behaviour of littoral fish. The present study investigates the effect of the time of day and the benthic habitat type on the shallow littoral fish community. Diel variations in total fish abundance, community composition, species abundance and behaviour were studied in two neighbouring stations in the Patraikos Gulf (Greece): one with cobble seabed cover and another with anthropogenic habitat structural modifications, like large artificial blocks, rocks and other objects. Visual census revealed that community composition differed depending on both the time of day and the habitat type. Total abundance was highest in the morning, whereas both diel abundance and habitat use were found to be taxon specific, with most species showing a preference for the artificial rocky habitat. Diel differences in species behavioural patterns were also observed, with a greater percentage of fish being active during the day (morning and noon) rather than in the afternoon. Although the fish community does not change entirely within a day or between adjacent locations, it is significantly variable in even small spatial and temporal scales. Consequently, there are implications for the design and implementation of sampling designs and monitoring plans that should be consistent in time of day throughout the sampling period and include locations structurally modified by humans. Furthermore, the management of the shallow littoral zone should consider the small-scale variability and alternate anthropogenic habitat with natural unaltered patches.

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. ​.

Trait-based life strategies, ecological niches, and niche overlap in the nekton of the data-poor Mediterranean Sea.

Biological traits can determine species ecological niches and define species responses to environmental variation. Species have a specific functional position in the biological community, resulting in interactions like interspecific competition. In this study, we used biological traits in order to define the life strategies of 205 nektonic species of the Mediterranean Sea. Furthermore, traits related to resource use were analyzed to determine the level of trait and niche overlap and their relationship to life strategies. Focusing on habitats of importance (Posidonia beds, coralligène formations, and lagoons), we investigated strategies and niches of the species present there. Finally, we examined the life strategy of Lessepsian species and investigated the niche overlap between them and indigenous species. Archetypal analysis indicated the existence of three life histories corresponding to strategies already documented for fish (equilibrium, periodic, and opportunistic), with some species also placed in intermediate positions. Niche overlap was evaluated by multiple correspondence analysis and the generation of a single distance metric between all species pairs. This identified species occupying relatively empty (underexploited) ecological niches, like the Lessepsian species Siganus luridus and S. rivulatus, a finding that can also be associated with their establishment in the Mediterranean. Most Lessepsian species were associated with the opportunistic life history strategy, again an important aspect related to their establishment. Also, we documented that most species occurring in important habitats have a relatively high overlap of niches. No significant differences were found in the life strategies across Mediterranean habitats; however, variation in niche overlap and traits related to habitat use was detected. The findings can be useful to determine theoretical competition between species and to identify empty ecological niches. Fisheries science can also benefit from comprehending the dynamics of competing stocks or predict the responses of data-poor stocks to anthropogenic stressors from known examples of species with shared life strategies.

Mediterranean nekton traits: distribution, relationships and significance for marine ecology monitoring and management.

Biological traits are increasingly used in order to study aspects of ecology as they are related to the organisms' fitness. Here we analyze a dataset of 23 traits regarding the life cycle, distribution, ecology and behavior of 235 nektonic species of the Mediterranean Sea in order to evaluate the distribution of traits, identify rare ones, detect relationships between trait pairs and identify species functional groups. Trait relationships were tested using correlation and non-linear regression for continuous traits, parametric and non-parametric inference tests for pairs of continuous-categorical traits and cooccurrence testing for categorical traits. The findings have significant implications concerning the potential effects of climate change (e.g., through the relationships of the trait of optimal temperature), fisheries or habitat loss (from the relationships of traits related to tolerance ranges). Furthermore, some unexpected relationships are documented, like the inversely proportional relationship between longevity and age at maturity as a percentage of life span. Associations between functional traits show affinities derived from phylogenetic constraints or life strategies; however, relationships among functional and ecological traits can indicate the potential environmental filtering that acts on functional traits. In total, 18 functional groups were identified by Hill-Smith ordination and hierarchical clustering and were characterized by their dominant traits. For the assessment of the results, we first evaluate the importance of each trait at the level of population, community, ecosystem and landscape and then propose the traits that should be monitored for the regulation and resilience of ecosystem functioning and the management of the marine ecosystems.

Evidence of subtle genetic structure in the sympatric species Mullus barbatus and Mullus surmuletus (Linnaeus, 1758) in the Mediterranean Sea.

Using thirteen microsatellite loci for Mullus barbatus and Mullus surmuletus collected in the Mediterranean Sea, the biogeographic boundaries, genetic distribution among and within basins and the impact of prolonged exploitation in both species were investigated as a basis for understanding their population dynamics and for improving Mullus spp. stock management. Different level of diversity indices among these co-occurring species were obtained, with M. barbatus showing higher allele richness and higher mean observed and expected heterozygosity than M. surmuletus. Reduced contemporary effective population size (Ne) and M-ratio values found in both species likely reflects recent demographic changes, due to a combination of high fishing pressures, habitat fragmentation and naturally occurring fluctuations in population size. Different patterns of genetic connectivity among populations sampled within the Mediterranean were observed for both species. Higher genetic structure was found for M. barbatus as opposed to a more homogenous pattern observed in M. surmuletus samples. Adriatic populations, previously considered panmictic and isolated from other Mediterranean regions, showed geographical partitioning within the basin but also population connectivity with the northern Ionian and Tyrrhenian Seas. Our results highlight the need for temporal sampling in understanding the complex pattern of population connectivity in the Mediterranean, particularly for management purposes.

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.