A coupled movement and bioenergetics model to explore the spawning migration of anchovy in the Bay of Biscay

Adult anchovies in the Bay of Biscay perform north to south migration from late winter to early summer for spawning. However, what triggers and drives the geographic shift of the population remains unclear and poorly understood. An individual-based fish model has been implemented to explore the potential mechanisms that control anchovy's movement routes toward its spawning habitats. To achieve this goal, two fish movement behaviors – gradient detection through restricted area search and kinesis – simulated fish response to its dynamic environment. A bioenergetics model was used to represent individual growth and reproduction along the fish trajectory. The environmental forcing (food, temperature) of the model was provided by a coupled physical–biogeochemical model. We followed a hypothesis-testing strategy to actualize a series of simulations using different cues and computational assumptions. The gradient detection behavior was found as the most suitable mechanism to recreate the observed shift of anchovy distribution under the combined effect of sea-surface temperature and zooplankton. In addition, our results suggested that southward movement occurred more actively from early April to middle May following favorably the spatio-temporal evolution of zooplankton and temperature. In terms of fish bioenergetics, individuals who ended up in the southern part of the bay presented better condition based on energy content, proposing the resulting energy gain as an ecological explanation for this migration. The kinesis approach resulted in a moderate performance, producing distribution pattern with the highest spread. Finally, model performance was not significantly affected by changes on the starting date, initial fish distribution and number of particles used in the simulations, whereas it was drastically influenced by the adopted cues.

Assessing natural mortality of Bay of Biscay anchovy from survey population and biomass estimates

La fermeture de la pêche de l’anchois (Engraulis encrasicolus) du Golfe de Gascogne entre 2005 et 2010, en raison de la faible biomasse du stock, a été une opportunité pour estimer la mortalité naturelle à partir de données de campagnes halieutiques issues de l’observation de la présence d’oeufs (DEPM) et de mesures acoustiques, utilisées pour l’évaluation de cette population depuis 1987. En considérant que la mortalité naturelle est constante au cours du temps et que la capturabilité des deux campagnes est identique pour tous les âges, la mortalité naturelle peut être estimée par des modèles log-linéaires appliquées aux séries temporelles de nombres aux âges issues des campagnes, et par des modèles d’évaluations de stock prenant en compte la saison. L’analyse suggère des valeurs de M autour de 0.9 comme mortalité naturelle courante à tous les âges. Cependant, nous avons des éléments forts indicateurs que la mortalité aux âges 2 et plus (M2+) est remarquablement plus forte qu’à l’âge 1 (M1) ce qui suppose un signe de mortalité sénescente, une possibilité qui a déjà été évoquée il y a longtemps pour ce type d’espèce à vie courte.

Validation of age determination using otoliths of the European anchovy (Engraulis encrasicolus L.) in the Bay of Biscay

Validation of the age determination procedure using otoliths of European anchovy in the Bay of Biscay was achieved by monitoring very strong year-classes in successive spring catches and surveys, as well as the seasonal occurrence of edge types. Historical corroboration of the ageing method was obtained by cross-correlation between successive age groups by year-classes in catches and surveys (1987–2013). Summary annual growth in length is also presented. Yearly annuli consist of a hyaline zone (either single or composite) and a wide opaque zone, disrupted occasionally by some typical checks (mainly at age-0 and age-1 at peak spawning time). Age determination, given a date of capture, requires knowledge of the typical annual growth pattern of otoliths, their seasonal edge formation by ages and the most typical checks. Most opaque growth occurs in summer and is minimal (translucent) in winter. Opaque zone formation begins earlier in younger fish (in spring), and this helps distinguish age-1 from age-2+.

Habitat selection and phenotypic diversity in the anchovy of the Bay of Biscay

Diversity among individuals in a population is an important feature linking vital rates with behaviour and spatial occupation. We measured the growth increments in the otolith of individual fishes collected on the annual fisheries survey PELGAS from 2001 to 2015. Individuals who grew larger at juvenile stage occupied later in life more off-shore habitats. Further, we analysed the allozymes of 13 different loci from 2001 to 2006. Alleles of the enzyme IDH showed different frequencies in inshore and offshore habitats. The population spatially segregates along a coast to off-shore gradient with individuals showing different early growth and allele frequencies. Results show how individuals in a population segregate spatially in different habitats in relation with phenotypic diversity. This implies modelling the population with individual-based and physiological approaches to fully grasp its dynamics. It also implies developing management strategies to conserve infra-population diversity as a means to garantee the occupation of the full range of habitats.

Influence of environmental variability and age on the body condition of small pelagic fish in the Gulf of Lions

Endogenous and environmental variables are fundamental in explaining variations in fish condition. Based on more than 20 yr of fish weight and length data, relative condition indices were computed for anchovy and sardine caught in the Gulf of Lions. Classification and regression trees (CART) were used to identify endogenous factors affecting fish condition, and to group years of similar condition. Both species showed a similar annual cycle with condition being minimal in February and maximal in July. CART identified 3 groups of years where the fish populations generally showed poor, average and good condition and within which condition differed between age classes but not according to sex. In particular, during the period of poor condition (mostly recent years), sardines older than 1 yr appeared to be more strongly affected than younger individuals. Time-series were analyzed using generalized linear models (GLMs) to examine the effects of oceanographic abiotic (temperature, Western Mediterranean Oscillation [WeMO] and Rhone outflow) and biotic (chlorophyll a and 6 plankton classes) factors on fish condition. The selected models explained 48 and 35% of the variance of anchovy and sardine condition, respectively. Sardine condition was negatively related to temperature but positively related to the WeMO and mesozooplankton and diatom concentrations. A positive effect of mesozooplankton and Rhone runoff on anchovy condition was detected. The importance of increasing temperatures and reduced water mixing in the NW Mediterranean Sea, affecting planktonic productivity and thus fish condition by bottom-up control processes, was highlighted by these results. Changes in plankton quality, quantity and phenology could lead to insufficient or inadequate food supply for both species.

Diversité phénotypique dans la population d’anchois commun (Engraulis encrasicolus) du golfe de Gascogne

With the constant rising of the human population, it is essential to assess and manage the exploitation of the marine species for human consumption such as the anchovy of Bay of Biscay. Thanks to the PELGAS annual surveys, stocks of each species are estimated (biomass, repartition) and studies are undertaken to improve our knowledge about these species. Allozymic studies realized on the anchovy allowed to find a correlation between 2 alleles of the locus IDH and the habitat (depth). This allozyme has a key role in Krebs’ cycle,being speed limiting.. It enables to tolerate variations of temperature and dissolved oxygen. It would be interesting to do new research on variations in the anchovy’s environment and its migrations.

Direct assessment of small pelagic fish by the PELGAS16 acoustic survey

The Pelgas16 acoustic survey has been carried out with very good weather conditions (regular low wind, medium temperatures) for the whole area, from the South of the bay of Biscay to the west of Brittany. The help of commercial vessels (two pairs of pelagic trawlers and a single one) during 18 days provided about 120 valid identification hauls instead of about 60 before 2007 when Thalassa was alone to identify echotraces. Their participation increased the precision of identification of echoes and some double hauls permitted to confirm that results provided by the two types of vessels (R/V and Fishing boats) were comparable and usable for biomass estimate purposes. These commercial vessels participated to the PELGAS survey in a very good spirit of collaboration, with the financial help of "France Filière Pêche" which is a groupment of French fishing organisations. The PELGAS16 survey observed a medium level of anchovy biomass, which seems to be a medium biomass compared to previous year's, comparable to 2010 and far away from the 2015 biomass. The biomass estimate of sardine observed during PELGAS16 is decreasing compared to the last years level of biomass. It confirms that this specie shows a variable abundance in the bay of Biscay at this period.

Analyse du compartiment mésozooplanctonique et écologie alimentaire printanière de la sardine, Sardina pilchardus (Walbaum, 1782), et de l’anchois, Engraulis encrasicolus (Linné, 1758) adultes dans le Golfe de Gascogne

Dietary studies of marine species constitute an important key to improve the understanding of its biology and of its role in the ecosystem. Thus, prey-predator relationships structure and determine population dynamics and the trophic network at the ecosystem scale. Among the major study sites, the marine ecosystem is submitted to natural and anthropogenic constraints. In the North-Eastern part of the Atlantic Ocean, the Bay of Biscay is a large open area surrounded South by Spain and East by France. This bay is an historic place of intense fishery activities for which the main small pelagic species targeted are the pilchard, Sardina pilchardus and the anchovy, Engraulis encrasicolus. The aim of this work is to analyze the trophic ecology of these two small pelagic fish in spring in the Bay of Biscay. To do this, a first section is devoted to their prey composed by the mesozooplanktonic compartment, through a two-fold approach: the characterization of their spatio-temporal dynamics during the decade 2003-2013 and the measurement of their energetic content in spring. For this season, it appears that all prey types are not worth energetically and that the Bay of Biscay represents a mosaic of dietary habitat. Moreover, the spring mesozooplankton community presents a strong spatial structuration, a temporal evolution marked by a major change in abundance and a control by the microphytoplankton biomass. The second section of this work is relative to a methodological approach of the trophic ecology of S. pilchardus and E. encrasicolus. Three different trophic tracers have been used: isotopic ratios of carbon and nitrogen, parasitological fauna and mercury contamination levels. To improve the use of the first of these trophic tracers, an experimental approach has been conducted with S. pilchardus to determine a trophic discrimination factor. Finally, it appears that the use of these three trophic tracers has always been permitted to highlight a temporal variability of the relative trophic ecology of these fish. However, no spatial dynamics could be identified through these three trophic tracers.

Body reserves mediate trade-offs between life-history traits: new insights from small pelagic fish reproduction

Limited resources in the environment prevent individuals from simultaneously maximizing all life-history traits, resulting in trade-offs. In particular, the cost of reproduction is well known to negatively affect energy investment in growth and maintenance. Here, we investigated these trade-offs during contrasting periods of high versus low fish size and body condition (before/after 2008) in the Gulf of Lions. Female reproductive allocation and performance in anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) were examined based on morphometric historical data from the 1970s and from 2003 to 2015. Additionally, potential maternal effects on egg quantity and quality were examined in 2014/2015. After 2008, the gonadosomatic index increased for sardine and remained steady for anchovy, while a strong decline in mean length at first maturity indicated earlier maturation for both species. Regarding maternal effects, for both species egg quantity was positively linked to fish size but not to fish lipid reserves, while the egg quality was positively related to lipid reserves. Atresia prevalence and intensity were rather low regardless of fish condition and size. Finally, estimations of total annual numbers of eggs spawned indicated a sharp decrease for sardine since 2008 but a slight increase for anchovy during the last 5 years. This study revealed a biased allocation towards reproduction in small pelagic fish when confronted with a really low body condition. This highlights that fish can maintain high reproductive investment potentially at the cost of other traits which might explain the present disappearance of old and large individuals in the Gulf of Lions.