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In this study we consider the applicability of Bergmann’s rule to the populations of small pelagic fish species in the Mediterranean Sea. Under Bergmann’s rule, body size increases with decreasing temperature and increasing latitude. Although this macroecological pattern in body sizes is well established for many taxa of endotherms and ectotherms, it does not remain universal, and the proposed mechanisms underlying it are multiple and still lack consensus. Here we explored the occurrence of geographical body size clines using measurements of average body sizes of 10 species collected in pelagic trawl hauls carried out during acoustic surveys in the Mediterranean Sea. Bergmann’s rule was evaluated by correlating body sizes with latitude and temperature for each species while accounting for potential confounding variables and sampling bias. For 5 species, namely anchovy, sardine, Atlantic chub mackerel, bogue and blue jack mackerel, we demonstrate that they follow Bergmann’s rule, with a decline in average body size by about 3.01, 3.43, 3.67, 3.82 and 3.76 % per 1°C of warming, respectively, although this did not translate with an increase in size with latitude. The adherence of these 5 pelagic fish to Bergmann’s rules strongly suggest that temperature is a major determinant of their body sizes and enables them to act as sentinel species for identifying the drivers and consequences of warming in the Mediterranean ecosystems

Size at first sexual maturity (L50) represents an important life-history trait that needs to be considered in the development of management measures as it provides fundamental information for avoiding the exploitation of younger individuals. L50 is known to display variability due to fishing pressure, geographical gradients, and environmental features. In this study, to investigate L50 variability among areas in the Mediterranean and Black Seas, maturity ogives of anchovies (Engraulis encrasicolus) were estimated by considering samples collected during the anchovy spawning period in the framework of the MEDiterranean International Acoustic Survey (MEDIAS) program. Anchovy size and sexual maturity data from several geographical subareas (GSAs), i.e., northern Spain, Gulf of Lion, Tyrrhenian Sea, Strait of Sicily, Adriatic Sea and Black Sea, were gathered according to a standard methodological protocol. Maturity ogives were estimated by a logistic regression considering total length, condition factor, sex and GSA. The obtained results showed a significant effect of the condition factor, in that fish in better condition reached maturity earlier, and the results also indicated differences in L50 values among the areas and between the sexes, with males reaching maturity at lower lengths than females. Even though the obtained L50 estimates are relative to the spawning period only, the variability observed at the Mediterranean basin scale highlights the importance of explicitly considering specific habitat characteristics when providing management advice based on an ecosystem approach for fisheries.

The study of small pelagic fish by hydroacoustic methods in the Mediterranean Sea began in the Adriatic in the 1950’s. Since 2009, internationally coordinated, extensive, regular surveys have been conducted in the framework of the MEDIAS - Mediterranean International Acoustic Surveys – action, under the EU Data Collection Framework, to provide inputs for the management of small pelagics, particularly European anchovy (Engraulis encrasicolus) and European sardine (Sardina pilchardus). The surveys cover EU Mediterranean waters and monitor the distribution and abundance of small pelagic species using a common protocol. The hydroacoustic surveys, historically conducted by each Country separately, have since been standardized and harmonized. This is a review of the evolution of the surveys from the beginning up to the present and discusses the future prospects. It reports the historical time series of anchovy, sardine and other small pelagics in different areas and describes the spatial and temporal distribution of small pelagic species in decadal maps of the major areas of their distribution in the European Mediterranean Sea. Furthermore, it discusses the use of MEDIAS data for stock assessment purposes, the collection of auxiliary information for an ecosystem-based management approach, the need for further standardization and future challenges.

Small pelagic fish play a key role in the marine ecosystem, controlling predator abundance and regulating primary production rates by foraging on plankton. Alterations induced at different ecological levels by fishing activities and/or environmental changes are affecting the reproductive success of several small pelagics, including European anchovy (Engraulis encrasicolus), which is a major target of Adriatic mid-water trawl and purse-seine fisheries. In this study, we evaluated short-term genetic changes of the species in the Adriatic Sea by applying molecular markers in samples of three generations of European anchovy. Thirteen polymorphic microsatellite markers and a mitochondrial gene were used in cohorts of adults and larvae, collected at three sites in the north-east, central, and southern Adriatic Sea in 2015. Furthermore, temporal, and spatial genetic variation was assessed by comparing the above dataset with a sample of adult anchovy collected in 2012 at three sites close to those sampled in 2015. Expected heterozygosity was higher in adults than in larvae, suggesting a loss of genetic diversity and uneven reproduction. In addition, a comparison of the two datasets demonstrated a change in the anchovy population structure from 2012 to 2015. In the reproductive event of 2015, this change led the two main genetic stocks described in the Adriatic Sea to merge into one. We suggest that the population structure of European anchovy in the north-eastern Adriatic may be influenced by changes in environmental parameters and by periodic alterations in the temporal pattern of population connectivity.

Brosset, Pablo ... et al.-- 14 pages, 7 figures, 2 tables, Small pelagic fish are among the most ecologically and economically important marine fish species and are characterized by large fluctuations all over the world. In the Mediterranean Sea, low catches and biomass of anchovies and sardines have been described in some areas during the last decade, resulting in important fisheries crises. Therefore, we studied anchovy and sardine body condition variability, a key index of population health and its response to environmental and anthropogenic changes. Wide temporal and spatial patterns were investigated by analyzing separately data from scientific surveys and fisheries in eight Mediterranean areas between 1975 and 2015. Results showed that anchovy and sardine body condition as well as maximum size in some areas sharply decreased in most Mediterranean areas along years (except in the Northern Alboran Sea). Despite this general pattern, well-marked environmental differences between sub-regions were highlighted by several analyses and variations in body condition were not found to be homogeneous over all the Mediterranean Sea. Further, other analyses revealed that except for the Adriatic where major changes towards a lower body condition were concomitant with a decrease in river runoffs and chl-a concentration, no concomitant environmental regime shift was detected in other areas. Together, these analyses highlighted the current poor body condition of almost all small pelagic fish populations in the Mediterranean. Yet, global environmental indices could not explain the observed changes and the general decrease in condition might more likely come from regional environmental and/or anthropogenic (fishing) effects. A prolonged state of poor fish body condition, together with an observed reduced size and early age-at-maturity may have strong ecological, economic and social consequences all around the Mediterranean Sea., PB acknowledges doctoral fellowship support from the French Ministère de l’Education Nationale, de la Recherche et de la Technologie

Natural/synthetic Endocrine Disrupting Chemicals (EDCs) may display estrogenic activity and a lower potency than 17β-estradiol. Nonetheless, their concentrations and additive effects can affect the endocrine system and reproductive processes related to the Hypothalamic-Pituitary-Gonadal (HPG) axis. Because of their persistence in both the environment and biological systems, they ultimately target multi-level predators, including humans. We detected presence and effects of xenobiotics on wild anchovy Engraulis encrasicolus in the Western Adriatic Sea. Twenty-one PCBs and five organochlorines were detected on the order of ng g-1; vitellogenin, vitellogenin receptor and genes encoding for the zona radiata proteins were evaluated in gonad and/or liver and found transcribed in male specimens; in addition, intersex was histologically identified in the 13% of testis. Our results have developed the understanding of the European anchovy's reproductive toxicological risk and our approach could assist the comprehension of the complex dynamics of commercially relevant Teleost species.

Our knowledge for the distribution of Atlantic mackerel (Scomber scombrus) in the Mediterranean Sea is limited and fragmented. In the current work habitat suitability modeling was applied to summer acoustic surveys data of Atlantic mackerel juveniles derived from the north part of the Mediterranean (i.e., acoustic data from the Gulf of Lions, pelagic trawls held during acoustic surveys in Spanish Mediterranean waters, south Adriatic Sea, Strait of Sicily, and North Aegean Sea) using generalized additive models (GAMs) along with satellite environmental and bathymetry data. Bathymetry along with sea surface temperature and circulation patterns, expressed through sea level anomaly and the zonal component of the absolute geostrophic velocity, were the environmental variables best to describe nursery grounds. The selected model was used to produce maps presenting the potential nursery grounds of Atlantic mackerel throughout the Mediterranean Sea as a measure of habitat adequacy. However, the assessed potential nursery grounds were generally marked as “occasional,” implying that although there are areas presenting high probability to encounter Atlantic mackerel, this picture can largely vary from year to year stressing the high susceptibility of the species to environmental conditions. In a further step and toward a spatial management perspective, we have estimated and visualized the overlap between Atlantic mackerel and anchovy/ sardine juvenile grounds throughout the basin. Results showed that although the degree of overlapping was generally low, not exceeding 15% in general, this varied at a regional level going up to 30%. The potential of the output of this work for management purposes like the implementation of spatially-explicit management tools is discussed, SI

It is well known that temporal fluctuations in small populations deeply influence evolutionary potential. Less well known is whether fluctuations can influence the evolutionary potentials of species with large census sizes. Here, we estimated genetic population parameters from as survey of polymorphic microsatellite DNA loci in archived otoliths from Adriatic European anchovy (Engraulis encrasicolus), a fish with large census sizes that supports numerous local fisheries. Stocks have fluctuated greatly over the past few decades, and the Adriatic fishery collapsed in 1987. Our results show a significant reduction of mean genetic parameters as a consequence of the population collapse. In addition, estimates of effective population size (N-e) are much smaller than those expected in a fishes with large population census sizes (N-c). Estimates of N-e indicate low effective population sizes, even before the population collapse. The ratio N-e/N-e ranged between 10(-6) and 10(-8), indicating a large discrepancy between the anchovy gene pool and population census size. Therefore, anchovy populations may be more vulnerable to fishery effort and environmental change than previously thought.

The sustained exploitation of marine populations requires an understanding of a species' adaptive seascape so that populations can track environmental changes from short- and long-term climate cycles and from human development. The analysis of the distributions of genetic markers among populations, together with correlates of life-history and environmental variability, can provide insights into the extent of adaptive variation. Here, we examined genetic variability among populations of mature European anchovies (n = 531) in the Adriatic (13 samples) and Tyrrhenian seas (2 samples) with neutral and putative non-neutral microsatellite loci. These genetic markers failed to confirm the occurrence of two anchovy species in the Adriatic Sea, as previously postulated. However, we found fine-scale population structure in the Adriatic, especially in northern areas, that was associated with four of the 13 environmental variables tested. Geographic gradients in sea temperature, salinity and dissolved oxygen appear to drive adaptive differences in spawning time and early larval development among populations. Resolving adaptive seascapes in Adriatic anchovies provides a means to understand mechanisms underpinning local adaptation and a basis for optimizing exploitation strategies for sustainable harvests.

Anchovy (Engraulis encrasicolus, L.) is one of the most important commercial species of the northern and central Adriatic Sea, as well as one of the most productive fisheries in the whole Mediterranean. In the Adriatic Sea the stock of anchovy is shared between Italy, Croatia and Slovenia. A joint stock assessment was carried out using catch data from all the fleets for the time interval 1975-2009. Analyses were performed using estimates of natural mortality at age obtained by means of two different methods and two population dynamics methods based on the analysis of catch-at-age data: Laurec-Sheperd virtual population analysis (VPA) and integrated catch-at-age (ICA), both tuned to acoustic estimates of abundance. Gislason’s estimates for natural mortality appeared to be more realistic and were thus preferred for short-lived species. The general trend of biomass and fishing mortality is similar for the two models, highlighting the major collapse of the stock in 1987. Nevertheless, ICA has enough flexibility to combine all the data available without adding too much complexity in comparison with a VPA approach and seems to perform better in terms of the spawning stock biomass/recruitment relationship and diagnostics (i.e. the retrospective pattern). For the stock status, the exploitation rate from ICA is higher than the suggested threshold of 0.4 proposed by Patterson for small pelagic species. La anchoa (Engraulis encrasicolus, L.) es una de las especies comerciales más importantes del mar Adriático central y septentrional, a la vez que es una de las actividades pesqueras más productivas en todo el Mediterráneo. La población de anchoa en el mar Adriático es compartida por Croacia, Italia y Eslovenia: una evaluación conjunta de la población fue realizada usando los datos de captura de todas las flotas para el periodo 1975-2009. Se efectuaron análisis usando estimaciones de la mortalidad natural por edad obtenidos mediante dos métodos diferentes y dos métodos de dinámica poblacional basados en el análisis de datos de captura por edad: Análisis de Población Virtual (VPA) de Laurec-Sheperd y Análisis de Captura Integrada por Edad (ICA), ambos acordados en las estimaciones acústicas de la abundancia. Las estimaciones de mortalidad natural de Gislason aparentaban ser más realísticas y por lo tanto daba preferencia a especies de vida corta. La tendencia general de la biomasa y de la mortalidad por pesca es similar en los dos modelos, destacando la fuerte disminución de la población en 1987. Sin embargo, ICA permite una flexibilidad suficiente para combinar todos los datos disponibles sin agregar excesivas complejidades con respecto al enfoque VPA y aparenta un mejor desempeño en términos de relación de SSB-R y en términos of diagnósticos (por ejemplo: el patrón retrospectivo). En relación al estado de la población, la tasa de explotación del ICA es más alta que el umbral sugerido de 0.4 propuesto por Patterson para las especies de pequeños pelágicos.

The paper provides an overview of the state of the small pelagic fish in the Adriatic Sea. The North Adriatic is reviewed for the 1976–1998 period and the South Adriatic for the 1987–1998 period. First the fluctuations in time and in space of the pelagic biomass, as a whole and per species, are presented. Then the dramatic collapse of anchovy stock and its apparent association with the decrease of surface temperature is discussed. Finally the changes of the anchovy and sardine catches are compared and analysed with respect to their abundances in the sea. The conclusion is that acoustic and satellite methods are able to provide a large amount of information on the variability in the pelagic populations; this is essential for an appropriate management of these resources. There is still a need, however, to refine the methods and to integrate this kind of information with further environmental data (including natural predators).

Fish sampling is a critical aspect of acoustic surveys, because it is directly related to the “transformation” of echo into species biomass and subsequently affects the accuracy of acoustic estimates. In the present study, we investigated the differences between day and night sampling in a) the catch composition through certain diversity indices and b) the length frequency distribution of anchovy and sardine using catch data of pelagic hauls collected from four different regions of the European Mediterranean waters. In addition, the possible bias in trawl efficiency due to sampling time and the possible error introduced in acoustic estimates were investigated. No statistically significant differences were found between day and night in any of the parameters examined. The results showed that a more flexible strategy can be adopted to reduce the duration and the cost of acoustic sampling for small pelagic species. The advantages and disadvantages of the two sampling strategies are discussed, Sí