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The Mediterranean Sea shows a trend of increasing temperature and decreasing productivity from the western to the eastern basin. In this work we investigate whether this trend is reflected in the cephalopod assemblages found throughout the Mediterranean. Data obtained with bottom trawl surveys carried out during the last 22 years by EU Mediterranean countries were used. In addition to analysing spatial differences in cephalopod assemblages, we also analysed putative temporal changes during the last two decades. For this purpose, the basin was spatially divided into bioregions, the trawling grounds were subdivided into depth strata, and the dataset was split into two time series of 11 years each. All analyses were done using PRIMER software. The species richness did not vary with the longitudinal gradient, though in most bioregions it showed a mild decrease with depth before plummeting in the deepest waters. Cluster analysis revealed four different bathymetric assemblages in all bioregions. Despite the contrasting conditions between basins and the claims of biodiversity loss, our study revealed that spatial and temporal differences during the last two decades were restricted to changes in the relative abundance of species from a common pool of species inhabiting the whole Mediterranean. El mar Mediterráneo muestra un patrón de aumento de la temperatura y disminución de la productividad de la cuenca occidental a la oriental. En este trabajo se investiga si este patrón se refleja en las comunidades de cefalópodos que habitan el Mediterráneo. Se utilizaron datos obtenidos en campañas de arrastre de fondo realizadas durante los últimos 22 años por la mayoría de países mediterráneos de la UE. Junto con el análisis de las diferencias espaciales en las comunidades de cefalópodos, también se analizaron cambios temporales durante las dos últimas décadas. Para ello, la cuenca se dividió espacialmente en diferentes bioregiones, mientras que el conjunto de datos se dividió en dos series temporales de 11 años cada una. Todos los análisis se realizaron utilizando el software PRIMER. La riqueza específica no varió con el gradiente longitudinal, aunque en la mayoría de las bioregiones mostró una leve disminución con la profundidad antes de desplomarse en el estrato más profundo. El análisis cluster reveló cuatro comunidades batimétricas diferentes en todas las bioregiones. A pesar de las contrastadas condiciones ambientales entre las cuencas y las afirmaciones de pérdida de biodiversidad, nuestro estudio reveló que las diferencias espaciales y temporales durante las dos últimas décadas se limitaron a cambios en la abundancia relativa de las especies a partir de un conjunto faunístico común que habita todo el Mediterráneo.

Nearly 75% of European fishing vessels belongs to small-scale fisheries (SSF). However, SSF have received little attention compared to industrial fisheries. In the Mediterranean Sea, where the SSF traditionally had a high socio-economic relevance, there is a widespread interest in securing its sustainable exploitation and viability. Here we analyze the SSF from Mallorca (Balearic Islands) using fishery statistics from the last 25 years (1990–2014). Although fleet size declined markedly (−55%), landings remained constant which might be related to different, not mutually exclusive explanations such as a decrease in unreported catches, stagnant and closed market or technological creeping. Multivariate analysis revealed eight different fishing tactics corresponding to the main target species. Aggregated, these eight species accounted for 52% and 71% of SSF landings weight and value, respectively. The fishery of these species is markedly seasonal and the landings of most of them showed important fluctuations but no clear trends. In contrast with the claims of SSF having lower impact on the natural resources than other fisheries, surplus production models revealed a generalized overexploitation of these target stocks, especially for the most high-valued species. Simulations with a bioeconomic model showed that fishing effort reductions of 38% would improve the health of fish stocks while increasing the economic profits to as much as 15% from current profits. If all stocks were kept below their MSY level, the reduction in fishing effort would have to be of 53% from current values. If the economic yield from the fishery was to be maximized (MEY), fishing effort would need to be reduced by 28%. Owing to important quantities of unreported catches in the most high-valued species, output values of the stock status indicators and the bioeconomic modelling should be taken with care beyond tracking trends in the fishery, SI

Merino, Gorka ... et. al.-- 14 pages, 9 figures, 1 table, 2 appendix, Mediterranean demersal fisheries are highly multispecific and many of their target stocks are overexploited. In addition, rocketing fuel costs and low market prices of traditionally high-value species are challenging the viability of fisheries. Here, based on the numeric results of a simulation model, we conclude that this situation can be remedied by reducing both fishing mortality and fishing costs. According to our model results, fishing effort reductions of 48-71% would improve the health of fish stocks while increasing the economic profits of Mallorca islands bottom trawl fishery to as much as 1.9. M€ (146% higher than current profits). If all fish stocks were exploited at their MSY (or below) level, the reduction in fishing effort would have to be of 71% from current values. If equilibrium profits from the fishery were to be maximized (MEY), fishing effort would need to be reduced by 48%. These results must be taken with caution due the many sources of uncertainty of our analysis. The modeling tools used to estimate these values are conditional to the adequate treatment of two sources of uncertainty that are particularly problematic in Mediterranean fisheries: insufficiently known recruitment variability and lack of periodic evaluations of the state of many species. Our results show that fishing effort reductions would produce economic yield gains after a period of transition. Further studies on the benefits of changing the size-selection pattern of fisheries, on better estimation of stock-recruitment relationships and on better quantifications of the contribution of secondary species to these fisheries, are expected to improve the scientific recommendations for Mediterranean demersal fisheries toward sustainability principles. © 2014 Elsevier B.V., The research leading to these results has received funding from the European Community's Seventh Framework Programme (FP7/2007–2013) under grant agreement MYFISH n° 289257

International audience; Cephalopod assemblages at the scale of the entire Mediterranean Sea were analysed using information from 2 decades of standardized scientific bottom trawl surveys. Western and eastern assemblages (6 yr of data) were compared using a combined approach of multivariate ordination techniques and non-linear regressions. These methods enabled us to distinguish assemblages and simultaneously analyse the influence of geographic, bathymetric and environmental (sea surface temperature and chlorophyll a concentration) gradients on observed community patterns. Despite few differences in species composition between sub-basins, the relative contribution of species differed. Bathymetry was the primary structural driver for the cephalopod communities of both basins, and contributed to 3 assemblages (shallow water, upper slope and middle slope). Winter temperature influenced community assemblages more strongly in the western than in the eastern basin, in contrast to a small but consistent winter productivity influence on community assemblages in both basins. Thus, the environmental parameters analysed did not cause an immediate change in cephalopod assemblages, but rather an effect lagged by several months. Differences in the relative importance of environmental drivers show that different processes operate in the 2 basins. These results demonstrate similarities and differences between Mediterranean basins regarding important cephalopod functional groups. This information should help integrative ecosystem management approaches currently used in fisheries and conservation management.

Aim Marine conservation often prioritizes measures based on ecosystem diversity, missing important information on the size and structure of communities such as that provided by body size-spectra (SS). While most studies have focussed on temporal or spatial patterns of SS, there is no research investigating their spatial heterogeneity, which is a key community trait revealing mechanisms of resilience. We aimed to comparatively determine the main patterns of variation of fish SS and their spatial heterogeneity across geographic and bathymetric gradients. Location Shelf and slope Atlantic and Mediterranean demersal ecosystems. Methods Using the information obtained from the experimental surveys, we calculated fish SS at four bathymetric strata and investigated the influence of the biomass of small-sized and large-sized fish on their temporal and spatial variation. We also assessed the functional relationship between the mean pattern and the spatial heterogeneity of SS as a characteristic structural trait of a community. Results The variation in SS demonstrates that Atlantic communities were geographically and bathymetrically more connected, while those in the Mediterranean were more fragmented. The relationship between mean SS and its spatial heterogeneity was a community fingerprint differentiating two contrasting pathways of community recovery between Atlantic and Mediterranean ecosystems. Atlantic communities were recovering because of the temporal increasing contribution of large fish and a ‘rescue effect’ of neighbouring areas, which have spatially homogenized size structure. Mediterranean communities were, by contrast, spatially clustered and highly dependent on small-sized fish variability that hampers the capacity of ecosystems to recover after a chronic degradation. Main conclusions Patterns of SS variation depend on the spatial complexity of communities, which contrasts between Atlantic and Mediterranean systems. Integrative analyses of size-based indicators (mean and variance) provide a complete assessment of the structural properties of communities and characterize the resilience mechanisms of recovery against perturbations under contrasting ecological contexts.

Cephalopods and elasmobranchs are important components of marine ecosystems, whereby knowing the ecological role they play in the structure and dynamics of trophic networks is paramount. With this aim, stomach contents and stable isotopes of the most abundant elasmobranch and cephalopod species (5 and 18 species, respectively) inhabiting deep-sea ecosystems from the western Mediterranean were analyzed. The predators investigated encompassed different taxonomic groups, such as rays and sharks within elasmobranchs, and squids, octopuses and cuttlefishes within cephalopods. Specifically, we investigated ontogenetic shifts in diet, feeding strategies and prey consumption, trophic structure and potential dietary overlap between and within both taxonomical groups. Stable isotope analysis revealed ontogenetic shifts in diet in three elasmobranch (rays and sharks) and two cephalopod (octopuses and squids) species. Isotopic data showed a contrasting food source gradient (δ13C), from pelagic (squids and cuttlefishes) to benthic (octopuses and elasmobranchs). Stomach data highlighted a great variety of trophic guilds which could be further aggregated into three broad categories: benthic, benthopelagic and pelagic feeders. The combination of both stomach content and stable isotope analyses revealed a clear food partitioning among species. Mesopelagic prey were found to be an important food resource for deep-sea elasmobranchs and cephalopods, which could be related to the strong oligotrophic conditions in the area. The observed differences in feeding strategies within cephalopods and elasmobranchs should be taken into account when defining functional groups in trophodynamic models from the western Mediterranean. Our results also revealed that cephalopods play a key role for the benthopelagic coupling, whereas demersal elasmobranchs contribute primarily to a one-way flux accumulating energy resources into deep-sea ecosystems., SI

The Mediterranean Sea is characterized by large scale gradients of temperature, productivity and salinity, in addition to pronounced mesoscale differences. Such a heterogeneous system is expected to shape the population dynamics of marine species. On the other hand, prevailing environmental and climatic conditions at whole basin scale may force spatially distant populations to fluctuate in synchrony. Cephalopods are excellent case studies to test these hypotheses owing to their high sensitivity to environmental conditions. Data of two cephalopod species with contrasting life histories (benthic octopus vs nectobenthic squid), obtained from scientific surveys carried out throughout the Mediterranean during the last 20 years were analyzed. The objectives of this study and the methods used to achieve them (in parentheses) were: (i) to investigate synchronies in spatially separated populations (decorrelation analysis); (ii) detect underlying common abundance trends over distant regions (dynamic factor analysis, DFA); and (iii) analyse putative influences of key environmental drivers such as productivity and sea surface temperature on the population dynamics at regional scale (general linear models, GLM). In accordance with their contrasting spatial mobility, the distance from where synchrony could no longer be detected (decorrelation scale) was higher in squid than in octopus (349 vs 217 km); for comparison, the maximum distance between locations was 2620 km. The DFA revealed a general increasing trend in the abundance of both species in most areas, which agrees with the already reported worldwide proliferation of cephalopods. DFA results also showed that population dynamics are more similar in the eastern than in the western Mediterranean basin. According to the GLM models, cephalopod populations were negatively affected by productivity, which would be explained by an increase of competition and predation by fishes. While warmer years coincided with declining octopus numbers, areas of high sea surface temperature showed higher densities of squid. Our results are relevant for regional fisheries management and demonstrate that the regionalisation objectives envisaged under the new Common Fishery Policy may not be adequate for Mediterranean cephalopod stocks., SI

Cephalopods are sensitive to environmental conditions due to their fast growth rates and short life cycles, being thus considered good ecological indicators. Here, we analyzed western Mediterranean cephalopod paralarvae communities during two consecutive summers (2004 and 2005) characterized by contrasting oceanographic conditions. Our main aim was to determine to what extent such contrasting scenarios affected the spatial distribution of the cephalopod early life stages and decipher the main environmental factors affecting each particular taxon. The contrasting oceanographic conditions during 2004 and 2005 included differences in the shape and location of a regional frontal system, which brought about differences in the sea circulation and the temperature regime. Such contrasting scenarios were reflected in the spatial distribution and the frequencies of appearance of both the whole community and the individual taxa. There was not any major factor affecting most taxa equally, which points to differences in species sensitivity to external conditions. Moreover, our results evidenced that paralarvae populations were more influenced by regional sea salinity and temperature than by mesoscale hydrographic features. Our study reveals that moderate changes in sea temperature have a major effect on the presence of paralarvae populations, which has profound implications in the context of the current global warming., SI

Fluctuations in marine populations occur both in terms of abundance and distributional ranges, and this has major implications for marine ecology and fisheries management. However, little is known about the variability in, and factors influencing, spatial dynamics in marine groups other than fish. Using time series data from trawl surveys conducted in the western Mediterranean Sea from 1994 to 2012, we analysed the variability in population distribution (latitude, longitude and depth) of 2 cephalopod species with contrasting life histories, viz. the nektobenthic squid Illex coindetii and the benthic octopus Eledone cirrhosa. We investigated the influence of demographic information (population density and mean individual size) together with environmental variables (including chlorophyll a concentration, runoff, precipitation, temperature and climate) to identify the main drivers shaping the cephalopod distributions in 4 shelf regions with contrasting oceanographic and geographic conditions. Marked inter-annual fluctuations were found in the distribution of the 2 species. In general, squid and octopus populations were affected by the same variables, but the effect of each variable depended on the species and study region. Different life strategies resulted in contrasting responses to the environment, with I. coindetii associated with variables contemporary to spring surveys. In contrast, E. cirrhosa was primarily affected by conditions during the previous winter. Vertical distribution was mainly explained by individual body length, while chlorophyll, temperature or local climate proved to be more important for geographical displacements., SI

According to their main life history traits, organisms can be arranged in a continuum from fast (species with small body size, short lifespan and high fecundity) to slow (species with opposite characteristics). Life history determines the responses of organisms to natural and anthropogenic factors, as slow species are expected to be more sensitive than fast species to perturbations. Owing to their contrasting traits, cephalopods and elasmobranchs are typical examples of fast and slow strategies, respectively. We investigated the responses of these two contrasting strategies to fishing exploitation and environmental conditions (temperature, productivity and depth) using generalized additive models. Our results confirmed the foreseen contrasting responses of cephalopods and elasmobranchs to natural (environment) and anthropogenic (harvesting) influences. Even though a priori foreseen, we did expect neither the clear-cut differential responses between groups nor the homogeneous sensitivity to the same factors within the two taxonomic groups. Apart from depth, which affected both groups equally, cephalopods and elasmobranchs were exclusively affected by environmental conditions and fishing exploitation, respectively. Owing to its short, annual cycle, cephalopods do not have overlapping generations and consequently lack the buffering effects conferred by different age classes observed in multi-aged species such as elasmobranchs. We suggest that cephalopods are sensitive to short-term perturbations, such as seasonal environmental changes, because they lack this buffering effect but they are in turn not influenced by continuous, long-term moderate disturbances such as fishing because of its high population growth and turnover. The contrary would apply to elasmobranchs, whose multi-aged population structure would buffer the seasonal environmental effects, but they would display strong responses to uninterrupted harvesting due to its low population resilience. Besides providing empirical evidence to the theoretically predicted contrasting responses of cephalopods and elasmobranchs to disturbances, our results are useful for the sustainable exploitation of these resources, SI

Cephalopod populations show wide temporal fluctuations in abundance, which have usually been investigated at inter-annual scales related to environmental variability. However, cephalopods are also strongly linked to seasonal environmental fluctuations owing to their short life cycles and single seasonal breeding. Therefore, population abundance critically depends on the success of breeding and recruitment from the previous year and the optimization of resources in a narrow period of favorable conditions. This adaptation of population dynamics is paramount in marked oligotrophic systems, such as the western Mediterranean Sea. We used monthly landings per unit effort (LPUE) to explore the spatio-temporal variability in seasonal patterns of three cephalopod species (Illex coindetii, Eledone cirrhosa and Octopus vulgaris). Common trends across the study area were characterized for each species. In all cases, seasonal patterns were geographically aggregated in relation to differences in local environment (i.e., primary production and surface hydro-climatology). Variability in the mean seasonal pattern over time was also investigated under contrasting environmental or population regimes. The mean seasonal trend was more pronounced in regimes of high-population densities, suggesting a density-dependent control that can modify the strength of the environmental forcing in the seasonal patterns. Our study also evidences a spatial synchrony in the seasonal fluctuations of LPUEs. Scales of synchrony ranged from 70 to 200 km, indicating a patchy-aggregated spatial pattern as a part of complex population structures in the western Mediterranean. Improving our understanding of seasonal dynamics of cephalopods across temporal and spatial scales may lead to improved forecasts and management strategies., SI

Accumulating evidence shows that fishing exploitation and environmental variables can synergistically affect the population dynamics of exploited populations. Here, we document an interaction between fishing impact and climate variability that triggered a synchronic response in the population fluctuations of six exploited species in the Mediterranean from 1965–2008. Throughout this period, the fishing activity experienced a sharp increase in fishing effort, which caused all stocks to shift from an early period of underexploitation to a later period of overexploitation. This change altered the population resilience of the stocks and brought about an increase in the sensitivity of its dynamics to climate variability. Landings increased exponentially when underexploited but displayed an oscillatory behaviour once overexploited. Climatic indices, related to the Mediterranean mesoscale hydrography and large-scale north Atlantic climatic variability, seemed to affect the species with broader age structure and longer lifespan, while the global-scale El Niño Southern Oscillation index (ENSO) positively influenced the population abundances of species with a narrow age structure and short lifespan. The species affected by ENSO preferentially inhabit the continental shelf, suggesting that Mediterranean shelf ecosystems are sensitive to the hydroclimatic variability linked to global climate. (Oxford University Press)

Populations of the same species can experience different responses to the environment throughout their distributional range as a result of spatial and temporal heterogeneity in habitat conditions. This highlights the importance of understanding the processes governing species distribution at local scales. However, research on species distribution often averages environmental covariates across large geographic areas, missing variability in population- environment interactions within geographically distinct regions. We used spatially explicit models to identify interactions between species and environmental, including chlorophyll a (Chla) and sea surface temperature (SST), and trophic (prey density) conditions, along with processes governing the distribution of two cephalopods with contrasting life-histories (octopus and squid) across the western Mediterranean Sea. This approach is relevant for cephalopods, since their population dynamics are especially sensitive to variations in habitat conditions and rarely stable in abundance and location. The regional distributions of the two cephalopod species matched two different trophic pathways present in the western Mediterranean Sea, associated with the Gulf of Lion upwelling and the Ebro river discharges respectively. The effects of the studied environmental and trophic conditions were spatially variant in both species, with usually stronger effects along their distributional boundaries. We identify areas where prey availability limited the abundance of cephalopod populations as well as contrasting effects of temperature in the warmest regions. Despite distributional patterns matching productive areas, a general negative effect of Chla on cephalopod densities suggests that competition pressure is common in the study area. Additionally, results highlight the importance of trophic interactions, beyond other common environmental factors, in shaping the distribution of cephalopod populations. Our study presents a valuable approach for understanding the spatially variant ecology of cephalopod populations, which is important for fisheries and ecosystem management., SI

© 2015 Inter-Research. The squid Loligo vulgaris (LV) and L. forbesii (LF) are 2 cephalopod species occurring in the Atlantic Ocean and the Mediterranean Sea. Bathymetric segregation allows the co-existence of both species, with LV preferentially inhabiting the shallow shelf and LF living on the shelf-break and upper slope grounds. In this paper, the feeding habits of LV and LF were studied for the first time in the Mediterranean, by means of stomach content analysis (1452 and 900 individuals of LV and LF, respectively). The main objective was to determine the diet of both species, analysing temporal and ontogenetic diet changes and inferring predator-prey interactions. Fish were by far the most important prey in both squid, followed by crustaceans and cephalopods. Prey composition revealed the bathymetric segregation of both species in the Mediterranean. Whereas LV preferentially consumed typical coastal species of sparids and gobiids, LF preyed on slope inhabitants such as myctophids and euphausiids. Ontogenetic shifts of diet occurred in both squid, but took place at contrasting sizes, suggesting that the factors triggering them might be speciesspecific. The diet of small-sized LV individuals was more dependent on bottom-living organisms than in large individuals, which preyed mainly on benthopelagic fish. During the main reproductive period in spring, LV increased the consumption of highly nutritive prey such as polychaetes (nereidids). Size-related differences in LF diet during the second half of the year indicated a deeper distribution of large individuals, preferentially preying on mesopelagic species and being thus involved in benthic-pelagic coupling.

Fuelled by the increasing importance of cephalopod fisheries in Europe, scientists and stakeholders have demanded their assessment and management. However, little has been done to improve the data collection under the EU Data Collection Framework (DCF) in order to analyse cephalopod populations. While the DCF allows member states to design flexible national sampling programmes, it establishes the minimum data requirements (MDR) each state is obliged to fulfil. This study was performed to investigate whether such MDR currently set by the DCF allow the application of depletion models (DMs) to assess European cephalopod stocks. Squid and cuttlefish fisheries from the western Mediterranean were used as a case study. This study sheds doubt on the suitability of the MDR to properly assess and manage cephalopod stocks by means of DMs. Owing to the high plasticity of life-history traits in cephalopod populations, biological parameters should be estimated during the actual depletion period of the fished stocks, rather than performing triennial sampling as established by the DCF. In order to accurately track the depletion event, the rapid growth rates of cephalopods implies that their populations should be monitored at shorter time scales (ideally weekly or biweekly) instead of quarterly as specified by the DCF. These measures would not require additional resources of the ongoing DCF but a redistribution of sampling efforts during the depletion period. Such changes in the sampling scheme could be designed and undertaken by the member states or directly integrated as requirements., This study was performed under the Data Collection Framework (cofunded by the EU and the Spanish Institute of Oceanography, IEO) and the CONFLICT project (CGL2008-958; funded by the Spanish Ministry of Science and Innovation). SK was financed by an IEO-FPI grant (2011/11)

Increasing human pressures and global environmental change may severely affect the diversity of species assemblages and associated ecosystem services. Despite the recent interest in phylogenetic and functional diversity, our knowledge on large spatio-temporal patterns of demersal fish diversity sampled by trawling remains still incomplete, notably in the Mediterranean Sea, one of the most threatened marine regions of the world. We investigated large spatio-temporal diversity patterns by analysing a dataset of 19,886 hauls from 10 to 800 m depth performed annually during the last two decades by standardised scientific bottom trawl field surveys across the Mediterranean Sea, within the MEDITS program. A multicomponent (eight diversity indices) and multi-scale (local assemblages, biogeographic regions to basins) approach indicates that only the two most traditional components (species richness and evenness) were sufficient to reflect patterns in taxonomic, phylogenetic or functional richness and divergence. We also put into question the use of widely computed indices that allow comparing directly taxonomic, phylogenetic and functional diversity within a unique mathematical framework. In addition, demersal fish assemblages sampled by trawl do not follow a continuous decreasing longitudinal/latitudinal diversity gradients (spatial effects explained up to 70.6% of deviance in regression tree and generalised linear models), for any of the indices and spatial scales analysed. Indeed, at both local and regional scales species richness was relatively high in the Iberian region, Malta, the Eastern Ionian and Aegean seas, meanwhile the Adriatic Sea and Cyprus showed a relatively low level. In contrast, evenness as well as taxonomic, phylogenetic and functional divergences did not show regional hotspots. All studied diversity components remained stable over the last two decades. Overall, our results highlight the need to use complementary diversity indices through different spatial scales when developing conservation strategies and defining delimitations for protected areas., SI

Human activities have substantially changed the world’s oceans in recent decades, altering marine food webs, habitats and biogeochemical processes. Cephalopods (squid, cuttlefish and octopuses) have a unique set of biological traits, including rapid growth, short lifespans and strong life-history plasticity, allowing them to adapt quickly to changing environmental conditions. There has been growing speculation that cephalopod populations are proliferating in response to a changing environment, a perception fuelled by increasing trends in cephalopod fisheries catch. To investigate long-term trends in cephalopod abundance, we assembled global time-series of cephalopod catch rates (catch per unit of fishing or sampling effort). We show that cephalopod populations have increased over the last six decades, a result that was remarkably consistent across a highly diverse set of cephalopod taxa. Positive trends were also evident for both fisheries-dependent and fisheries-independent time-series, suggesting that trends are not solely due to factors associated with developing fisheries. Our results suggest that large-scale, directional processes, common to a range of coastal and oceanic environments, are responsible. This study presents the first evidence that cephalopod populations have increased globally, indicating that these ecologically and commercially important invertebrates may have benefited from a changing ocean environment, SI

Special issue The wrapping up of the IDEADOS project: International Workshop on Environment, Ecosystems and Demersal Resources, and Fisheries.-- 11 pages, 3 figures, 3 tables.-- Corrigendum to “Structure and dynamics of food webs in the water column on shelf and slope grounds of the western Mediterranean” [J. Mar. Syst. 138 (2014) 171–181], Journal of Marine Systems 139: 505–508 (2014) https://doi.org/10.1016/j.jmarsys.2014.09.005, Benthic-pelagic coupling is an important process connecting species throughout the water column, particularly, in deep-sea systems where faunal assemblages can be dense if indirectly sustained by production from the above. Through stable isotope analyses, this study explored the sources of production, trophic structure, and bentho-pelagic coupling in two locations with contrasting oceanographic conditions from the western Mediterranean, in the Balearic (BsB) and the Algerian (AsB) sub-basins. The samples of 89 dominant species (23 decapods, 19 cephalopods, 33 fishes, among the other taxa), inhabiting the hyperbenthic and pelagic domains, from the shelf break (250m), upper slope (650m), and middle slope (850m) were analyzed. Results suggested long food webs of approximately four trophic levels (TrLs) that were sustained by planktonic source material in shallower waters and degraded particulate organic matter of planktonic origin in deeper waters. Most of the collected species (70%) occupied intermediate trophic positions between the 3rd and 4th TrLs. The species δ15N and δ13C values exhibited a broad range, consistent with the high diversity that might be attributed to the oligotrophic conditions. As the depth increased, stronger segregation occurred between the trophic groups, and spatial differences were found among consumers of the two locations. Species in the AsB always had consistently higher δ15N values than in the BsB, which could possibly be attributed to the basal δ15N that was present through the food web. Despite the contrasting basin characteristics, a similarly close bentho-pelagic coupling pattern was observed at both locations, except at the deepest ground, especially at the AsB, where the mean δ13C values from the hyperbenthic and pelagic compartments were more distant. This could be related to the higher degree of reworking of organic matter in the AsB. Overall, these findings suggested the need for a depth-stratified approach to analyze deep-sea food webs in the study site in future studies. © 2014 Elsevier B.V., This research was supported by the project CTM2008-04489-C03-01 and CTM2008-04489-C03-02. The isotope analyses were funded by the Government of the Balearic Islands, Acció especial: AAEE0138/09 (DireccióGeneral de Recerca, Desenvolupament Tecnològic i Innovació, Conselleria d'Economia Hisenda i Innovació)

The cuttlefish Sepia officinalis constitutes an important fishery resource in the Mediterranean, where it is exploited by both the bottom trawl and small-scale fleet. However, there is currently scarce information on the Mediterranean stocks, since most studies on the population dynamics of this species have been undertaken in the northeast Atlantic. In this work we first analysed different aspects of the cuttlefish life-history from the western Mediterranean such as population structure, reproduction and the trade-offs between somatic condition and reproduction investments. Secondly, we investigated the effects of different environmental parameters (e.g. climate indices, sea surface temperature (SST), rainfall, chlorophyll-a concentration (Chla) and moon phase) on these populations, analysing several landing time series spanning the last 45 years. Our results revealed that Mediterranean cuttlefish populations exhibit strong seasonal variations owing to a reproductive migration towards coastal waters. The positive relationships between somatic and reproductive condition pointed to an income breeder strategy; this was reinforced by the percentage of empty stomachs, which was lowest just before the reproductive period peak. Despite the putative high sensitivity of cephalopod populations to external abiotic factors, our results showed that Mediterranean cuttlefish populations were not affected by most of the environmental parameters investigated. Significant effects were found for SST and a local climatic index, but no or very weak influences were evident for other parameters such as large-scale climatic phenomena (e.g. North Atlantic Oscillation, Mediterranean Oscillation) or other locally-related variables (e.g. rainfall, Chla). Our results revealed a shift in the cuttlefish population dynamics in the early 1980s, which could be related to important changes in the local hydroclimatology reported by previous authors.

Several aspects of the biology of Bathypolypus sponsalis were studied from 297 individuals (115 males, 180 females and 2 indeterminates) caught in a depth range of 200–800 m depth in the western Mediterranean Sea. The paper presents data on sizes (length-weight relationships, size-frequency distributions) and reproduction (sex ratio, maturation, condition), and also analyses of the diet of B. sponsalis from samples taken throughout the year. Length-weight relationships showed that females are heavier than males at the same mantle length. Although mature individuals were found all year round, the maximum number occurred in spring and summer. Sexual maturation data revealed that males mature at smaller sizes than females. The gonadosomatic index increased with maturity in both sexes; the increase was gradual in males, but abrupt in females. The digestive gland index was used as a condition index and showed a differential behaviour with maturity; it increased gradually in females, but decreased in males. Like other octopus species, B. sponsalis appears to be an opportunistic predator, feeding on a great variety of preys. Stomach content analysis yielded a total of 19 different prey items belonging to four major groups (Crustacea, Mollusca, Ophiuroidea and Osteichthya). The first three groups were the more frequent preys, since crustaceans, molluscs and ophiuroids appeared in 76%, 49% and 30% of the stomachs, respectively. Decapoda Reptantia (among crustaceans) and cephalopods and bivalves (among molluscs) constituted the more abundant prey items. While the Decapoda Reptantia group was significantly more abundant in stomach contents of females, gastropods were taken more frequently by males. These differences in diet could reveal females as a more active predators than males, Sí

The analysis of 79 hauls performed by commercial bottom trawlers from 50 to 800 m depth in the Balearic Sea (north-western Mediterranean) from June 1995 to September 1996 yielded a total of 30 cephalopod species belonging to 12 families. Cluster analysis of these data gave as a result two main groups 50–200 m and 200–800 m each subdivided into two other groups (50–100 vs. 100–200 m and 200–600 vs. 600–800 m). These results suggested the existence of two assemblages that could be associated to the continental shelf (50–100 m) and the upper slope (600–800 m) separated by a wide transitional zone (100–600 m) representing a region of overlapping shelf and slope faunas (ecotone). The faunistic bathymetric gradient showed a continuous substitution of species with depth rather than discrete assemblages separated by distinct boundaries. The more coastal species such as Eledone moschata, Loligo vulgarisSepia officinalis and Octopus vulgaris were found on the continental shelf; on the transitional zone, apart from species characteristic of this zone (Illex coindetii, Sepietta oweniana, Rossia macrosoma, Scaeurgus unicirrhus and Pteroctopus tetracirrhus), we also observed species from both the continental shelf and slope. The upper slope was characterized by typical species of deeper waters, such asBathypolypus sponsalis , Histioteuthis reversa, H. bonnellii, Ancistroteuthis lichtensteinii andOnychoteuthis banksii . The octopod O. vulgaris was the dominant species on the continental shelf and upper transitional zone, being substituted by T. sagittatus on the lower transitional zone and upper slope. Mean biomass decreased abruptly from the continental shelf to the transitional zone and from there to the upper slope. Mean species richness and species diversity were higher in the transitional zone than in the continental shelf and upper slope. Finally, some biological aspects of the more abundant deep-sea cephalopod species are studied: Bathypolypus sponsalis, Octopus salutii,Pteroctopus tetracirrhus , Histioteuthis reversa and H. bonnellii., Sí

Species composition, size range and some behavioral notes about drifting flotsam associated fish of the Balearic Islands are presented, and more detailed observations on Seriola dumerili, Naucrates ductor, Coryphaena hippurus and Scomberesox saurus are also given. 25 fish species belonging to 18 families were identified, 12 of these species and the individuals of the family Mugilidae were recorded for the first time associated with floating objects in the western Mediterranean. Most of the specimens collected were juveniles, which suggests the importance of floating objects as a nursery, and thus in the recruitment and redistribution of fishes. Analisis of two drift floating material samples show that objects of anthropogenic origin were most abundant (83.5% and 63.5%) and suggests that at present, human refuse may have taken over the role of the floating remains of marine plants for fishes in the western Mediterranean pelagic environment., No disponible

Stomach contents of Todarodes sagittatus caught by trawlers working from 100 to 800 m depth in the Balearic Sea (western Mediterranean) were studied. From the 348 stomachs examined (153 males and 195 females) 33.62% were empty (39.21% in males and 29.74% in females). The diet of the squid was composed of 58 different prey items belonging to four major groups: Osteichthya, Crustacea, Cephalopoda and Chondrichthya. Osteichthyes, crustaceans and cephalopods were the most common prey, with a frequency of occurrence value of 84.85, 48.92 and 29.87% respectively. A change in the diet as the squid grows was observed, since juveniles feed basically on fishes while adults prey more actively on crustaceans. Analysis of the diet by size-classes reflected an ontogenetic migration to deeper waters since, parallel to the increase of size, a raise in the percentage of prey species inhabiting deeper waters was detected. Cannibalism was quite frequent, since T. sagittatus was the second most common cephalopod prey. Females had higher fullness-weight index and lower emptiness index than males, which reflects their major energetic demand for egg production.

6 pages, 4 figures, 1 table, The aim of the study was to define the commercial importance of cephalopods caught by trawlers from five Italian and Spanish Mediterranean ports. From summer 1995 to spring 1996, samples of the catches of trawlers were taken on board. The type of fishing activity was recorded and data were collected from at least three commercial hauls for each port, season and bathymetric stratum (350 m). The species composition and yield of the retained and discarded portions of the catch were noted for each port. From these data the commercial importance of each species was evaluated. Cephalopods were commercially important particularly in the shallower bathymetric stratum (

The capture of a specimen of the oceanic cephalopod Taningia danae Joubin, 1931, caught during a bottom trawl survey carried out off the Algerian coast, is reported for the first time in the Mediterranean Sea. The individual, which was taken at 385-395 m depth, was a juvenile with a 55.6 mm mantle length and age of 132 days. This finding is important as there are few worldwide records of this species.

The stock concept plays a pivotal role in fisheries assessment and management. Stocks are defined according to biological, geographical, economic or socio-political factors. The General Fisheries Commission for the Mediterranean (GFCM) has established thirty management geographical sub-areas (GSAs) based on political and statistical considerations rather than biological or economic factors. Here, we present our view on the main biological and ecological aspects that should be considered for delineating different management units in the Mediterranean. We focus on the Balearic Islands (GSA05) as a case study highlighting its specificities compared to the adjacent coast of the Iberian Peninsula (GSA06), but the approach could be generalized to the problem of identifying stock boundaries in other areas. The work is based on published information from different marine disciplines such as geomorphology, ecology and fisheries, combined with the analysis of new data coming from official fishery statistics and scientific surveys. This approach avoids the important draw¬backs (inconclusive results, high costs) of other time-consuming techniques used in stock identification, such as genetics. According to the information presented, we conclude that GSA05 should be maintained as an individualized area for assess-ment and management purposes in the western Mediterranean, Sí

We assessed the elasmobranch communities exploited by the bottom trawl fishery off the Balearic Islands (northwestern Mediterranean) by analysing two time series containing information with different levels of detail: 1) a long-term series containing the abundance indices of the whole elasmobranch community landed from 1965 to 2009; and 2) a shorter time series from 2000 to 2009 from two different data sources (daily sales records: 2000-2009; bottom trawl surveys: 2001-2009), with information that allowed the shelf and slope to be analysed separately and different community parameters and abundance indices to be studied. Temporal trends of all these variables were used as ecological status indicators and tested with generalized additive models and analysis of variance. They were finally summarised using the Traffic Light methodology. The long-term series showed two marked trends, an inter-annual decreasing trend that would be a consequence of the large increase in fishing effort, and an intra-annual trend with a clear peak in May in accordance with the seasonal biological cycle of these species. However, data from the last ten years revealed differences between the shelf and slope, as some parameters increased significantly over time on the shelf but did not follow any trend on the slope. The Traffic Light methodology captured these differences well, demonstrating its usefulness for interpreting ecological transitions at a glance. Se han evaluado las poblaciones de elasmobranquios explotadas por la pesca de arrastre de las Islas Baleares (Mediterráneo noroccidental) analizando dos series temporales con información a distinto nivel de detalle: 1) una serie larga que contiene únicamente el índice de abundancia de toda la población de elasmobranquios desembarcada entre 1965 y 2009 y 2) una serie corta a lo largo de la primera década del siglo XXI proveniente de dos fuentes de datos diferentes (hojas de venta diarias: 2000-2009; campañas de arrastre de fondo: 2001-2009), que permitieron el análisis, para la plataforma y el talud por separado, de diferentes parámetros de la comunidad además de índices de abundancia. Las tendencias temporales de todas estas variables se usaron como indicadores ecológicos de estado y se testaron aplicando modelos generales aditivos y análisis de la varianza y finalmente se han resumido usando la metodología de los Semáforos. La serie larga mostró dos señales claras, un decrecimiento interanual que podría ser consecuencia del importante incremento del esfuerzo pesquero, y una tendencia intra-anual con un claro pico en mayo, relacionado con el ciclo biológico estacional de estas especies. Así y todo, los datos del primer decenio del siglo XXI revelaron diferencias entre la plataforma y el talud, porque algunos parámetros aumentaron significativamente con el tiempo en el primer caso, pero no siguieron ninguna tendencia en el segundo. Estas diferencias quedaron bien reflejadas en los Semáforos, demostrando su utilidad en reflejar las transiciones ecológicas.

In this paper we compare the elasmobranch composition, depth-related trends of species and community descriptors (abundance, biomass, mean fish weight, species richness and diversity) of the Balearic Islands Archipelago and Algeria (north- and southwestern Mediterranean, respectively). The samples used in this study were collected in bottom trawl surveys between 36 and 779 m depth. Generalized linear models, linear regression, cluster and similarity percentage analyses, and generalized additive models were used to compare the two areas. Twenty-nine elasmobranch species were caught, 12 of them common to both areas, 7 appearing only in the Balearic Islands, and 10 only in Algeria. The bathymetric distributions of species were, in most cases, best fitted to uni-modal response curves. The models for species common to both areas showed similar bathymetric trends, except for Etmopterus spinax, which in Algeria showed a maximum abundance lo¬cated 154 m shallower than in the Balearic Islands. Four bathymetric assemblages were identified in both areas with similar depth ranges but with different species composition. The mean values of the community descriptors were higher on the shelf in the Balearic Islands, whereas higher values were detected on the slope in Algeria. Different between-area bathymetric trends for all the descriptors were detected, with the exception of mean fish weight. The distinct environmental scenarios and fishing histories of the areas studied are discussed as underlying traits influencing the elasmobranch populations, Sí

Linking the abiotic and biotic traits of ecosystems is a critical step towards understanding their structure and functioning. Here we attempt to determine the connections between the hydrodynamics, benthic landscape and the associated fish communities on the coastal continental shelf off the Balearic Islands (western Mediterranean). Specifically we investigate the role the hydrodynamics play in shaping the benthic landscape, and whether the hydrodynamics affect the composition and structure of demersal fish communities. A realistic numerical model was used to establish the hydrodynamic characteristics of the area. The study area showed high hydrodynamic variability on a medium spatial scale (tens of km) in terms of mean water velocity (Ū). Principal component analysis was used to determine the main gradients of macro-epibenthic variability. Redundancy analysis (RDA) was used to model the effect of the hydrodynamics on macro-epibenthic species. RDA was also used to model the effect of the hydrodynamics and macro-epibenthos on the abundance of the associated fish fauna, and on its biomass at a community level using biomass spectra classes. The results showed that the hydrodynamics had a significant influence on the distribution of both macro-epibenthic species and the associated fish species. The latter was also influenced by the macro-epibenthos. Fish size appeared to be a key attribute for the distribution of species across gradients of Ū and macro-epibenthic change. Our findings can be applied in ecosystem-based fisheries management, as they show that it is necessary to take into account both the biotic and abiotic traits of the habitats when the habitat use and requirements of the associated species are defined. © 2011 Elsevier Ltd., This paper is a result of the MIGJORN project co-financed by the Conselleria d'Agricultura i Pesca of the Govern de les Illes Balears and the Spanish Institute of Oceanography; and the INDEMARES project (EU Life project) and Fundación Biodiversidad.

The vast majority of modern cephalopods is single-season breeders, or semelparous in the strict sense, that die soon after the reproduction takes place. Individual body condition in these marine invertebrates is expected to be highly affected by reproduction because: 1) the gonad weight of females, which represents

Risso's dolphins, pilot whales and sperm whales rarely strand in the northwestern Mediterranean. Thus, their feeding ecology, through the analysis of stomach contents, is poorly known. The aim of this study was to gain further insight into the segregation/superposition of the diet and habitat of Risso's dolphins, pilot whales and sperm whales using chemical tracers, namely, stable isotopes (d13C, d15N) and organochlorines. Significantly different d15N values were obtained in Risso's dolphins (11.7 +/- 0.7%), sperm whales (10.8 +/- 0.3%) and pilot whales (9.8 +/- 0.3%), revealing different trophic levels. These differences are presumably due to various proportions of Histioteuthidae cephalopods in each toothed whale's diet. Similar d13C contents between species indicated long-term habitat superposition or corroborated important seasonal migrations. Lower congener 180 concentrations (8.20 vs. 21.73 µg.g-1 lw) and higher tDDT/tPCB ratios (0.93 vs. 0.42) were observed in sperm whales compared with Risso's dolphins and may indicate wider migrations for the former. Therefore, competition between these species seems to depend on different trophic levels and migration patterns.

The life cycle of the two species of the deep-sea family Histioteuthidae inhabiting the Mediterranean Sea ( Histioteuthis reversa and Histioteuthis bonnellii ) was studied from monthly samples taken throughout the year during daytime hours by bottom trawl gears. A small sample of individuals found floating dead on the sea surface was also analyzed. Both species were caught exclusively on the upper slope at depths greater than 300 m. Their frequency of occurrence increased with depth and showed two different peaks, at 500–600 m and 600–700 m depth in H. bonnellii and H. reversa, respectively, which might indicate spatial segregation. Maturity stages were assigned using macroscopic determination and confirmed with histological analyses. Although mature males were caught all year round, no mature females were found, which suggests that their sexual maturation in the western Mediterranean takes place deeper than the maximum depth sampled (800 m). In fact, the increase in mean squid size with increasing depth in H. reversa indicates an ontogenetic migration to deeper waters. The individuals of both species found floating dead on the sea surface were spent females which had a relatively large cluster of small atresic eggs and a small number of remaining mature eggs scattered in the ovary and mantle cavity. The sizes of these females were clearly larger than the largest individuals caught with bottom trawls. A total of 12 and 7 different types of prey, belonging to three major taxonomic groups (crustaceans, osteichthyes and cephalopods), were identified in the stomach contents of H. reversa and H. bonnellii, respectively. In both species fishes were by far the main prey followed by crustaceans, whereas cephalopods were found only occasionally. The preys identified, mainly myctophids and natantian crustaceans, indicate that both histioteuthids base their diet on pelagic nictemeral migrators.

Scorpaena notata is a small, sedentary scorpaenid species widely distributed in the Mediterranean and adjacent waters of the Atlantic. In the western Mediterranean it inhabits coastal continental shelf bottoms. In the Balearic Islands, these bottoms are characterised by the presence of the facies with red algae, including both Peyssonnelia and mäerl beds. These beds enhance the structural complexity, biodiversity and secondary production of the soft bottoms. Due to the oceanographic conditions of the Islands, the facies with red algae are especially rich in terms of biomass and algal coverage, and are widespread distributed between 40 and 90 m depth, where trawlers exploiting the continental shelf operate. The present work studies the biology of S. notata and its relationship with habitat characteristics. Special attention is focused on the aspects related to fish condition and growth as a tool to assess the importance of the facies with red algae for fish. The reproduction period of S. notata in the Balearic Islands occurs in summer and is accompanied by a decrease in hepatic condition, as it happens in the adjacent area off the Iberian Peninsula; however, in contrast with this adjacent area, this period is accompanied by a decrease in somatic condition and an increase in feeding potential, which suggests that these could be adaptations to the higher oligotrophy of the Archipelago. The standardised algal biomass (mostly Rhodophyceae) present in the bottoms positively affected the abundance, somatic condition and feeding potential of S. notata. Individuals inhabiting bottoms with the highest algal biomass showed faster growth than the entire population analysed together. Both, the structural complexity and the availability of preys in the facies with red algae are revealed as advantageous traits for the life history of fish. Taking into account the importance of individual health for the overall success of the population, the indexes studied here could be a useful tool for identifying high quality or essential fish habitats. Our results highlight the importance of the facies with red algae as oasis of high productivity where benthic fish can circumvent the general oligotrophic conditions of the Mediterranean, and the necessity of urgent management measures in order to protect them from human impacts., Sí

The spatial and temporal variations of deep-sea megafaunal assemblages from the western Mediterranean are analysed in the present paper. The assemblages from two locations of the Balearic Islands situated 120 km apart were compared using data collected seasonally on a bathymetric stratum covering the 150–750 m depth range during six bottom-trawl surveys. The assemblage structure, in terms of species composition, species dominance and population sizes, was differentially affected by the spatio-temporal variables analysed (depth, location and fishing period). Although depth was the main factor determining the assemblage composition, the differences obtained between the two locations were also relevant. On the upper slope these between-location differences in the dynamics of megafaunal assemblages were found to be related to the effect of fishing exploitation. Population size-based metrics and biomass spectra were good predictors of meso-scale fishing effects, and were mainly reflected by elasmobranchs and demersal teleosts. Nevertheless, the effects of fishing depended on the species considered. Two dominant large-sized fish species found on the upper slope in both localities, Galeus melastomus and Phycis blennoides , had higher biomass values associated with lower fishing effort. Although the mean body weight (MBW) of both species and also the mean maximum body weight (MMBW) of G. melastomus agreed with this pattern, the P. blennoides MMBW did not. This last case could be indicative of natural size-trends such as the bigger–deeper phenomenon which refers to the displacement of large individuals towards the deeper limit of their bathymetric distribution, beyond the maximum depth sampled in this study for this species. By contrast, the target species of the upper slope fishery, the red shrimp Aristeus antennatus, was not negatively affected by the direct impact of fishing activity and other environmental factors, such as the presence of specific water masses could also be important.

The capture of a specimen of the oceanic cephalopod Taningia danae Joubin, 1931, caught during a bottom trawl survey carried out off the Algerian coast, is reported for the first time in the Mediterranean Sea. The individual, which was taken at 385-395 m depth, was a juvenile with a 55.6 mm mantle length and age of 132 days. This finding is important as there are few worldwide records of this species, Sí

We investigate the effects of the Columbretes Islands Marine Reserve (CIMR, Western Mediterranean) on the adjacent Palinurus elephas (Fabricius, 1787) fishery. After 9 to 12 yr of no-take protection there was a gradient of lobster density from the interior of the reserve up to a distance of about 4 km from its boundary. Catch and effort data were collected onboard commercial fishing boats in the fishery adjacent to the CIMR, and combined with catch per unit effort (CPUE) data from monitoring surveys conducted annually inside the reserve. Generalized additive (GAM) and linear (GLM) models were employed to examine the relationships of CPUE and catch per unit area (CPUA) as a function of distance to the reserve boundary. CPUE showed a significant non-linear decline with distance from the centre of the reserve, with a depression at the boundary followed by a plateau. This depression was caused by local depletion associated with concentration of fishing effort at the reserve boundary, while the plateau suggests that lobster export from the reserve is sufficient to maintain stable catch rates up to 1500 m from the boundary. Commercial catch and effort data were combined to estimate CPUA, which declined linearly with distance from the reserve. Analysis of recaptures of lobsters tagged and released inside the reserve indicates that the density gradient is caused by lobsters emigrating from the reserve, Sí

A total of 310 individuals (124 males, 185 females, 1 indeterminate) of the octopod Octopus salutii caught throughout the year by bottom trawlers from 25 to 800 m depth in the western Mediterranean were analysed to study some general aspects of its biology. Octopus sizes ranged from 4.0 to 13.0 cm ML and 3.5 to 16.5 cm ML in males and females, respectively. The species inhabits the lower continental shelf and upper slope, primarily between 250 and 500 m depth. The scarcity of small-sized individuals in the samples and the predominance of males and females >8 and >9 cm ML, respectively, suggest two interpretations: (1) juveniles inhabit grounds inaccessible to, or unsampled by, trawlers, or they avoid capture owing to behavioural aspects; or (2) the species meet in the trawling grounds for reproduction, because in the sizes described the majority of individuals have begun sexual maturation. The reproductive period was found to occur during spring and summer. The stomach content analysis revealed a diet composed of 33 different prey items belonging to three major taxonomic groups (crustaceans, fishes and cephalopods). Quantitatively, crustaceans (Decapoda Reptantia and Natantia groups) were the most important prey, appearing in 87% of the stomachs, followed by fish (25%) and cephalopods (10%). Finally, the paper presents data on mantle length–total weight, gonad weight–total weight and ligulae length–total weight relationships, and utilises different indexes (gonadosomatic, hepatosomatic, fullness weight and emptiness) to describe changes through the three maturity stages of both sexes, Sí