Influence of environmental factors on different life stages of European anchovy (Engraulis encrasicolus) and European sardine (Sardina pilchardus) from the Mediterranean Sea: A literature review

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Highlights

  • We reviewed the literature available on how environmental factors affect anchovy and sardine.

  • The studies were heterogeneously distributed.

  • The most studied variables were temperature, depth, salinity and Chl-a.

  • Temperature was one of the most influencing variables for both species.

  • More information on the environmental effect on larvae and egg is needed.

Abstract

European anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) are two of the most important small pelagic fish species in terms of biomass and commercial interest in the Mediterranean Sea. Their populations are clearly influenced by environmental variations, making them good bio-indicators of climate-driven changes. Here, we reviewed the different environmental factors documented to affect anchovy and sardine biology by using a systematic review approach on methodologies used to study environmental effects on these species, variables identified to play a role, and relationships found between ecological variables and the environment. Our results show that available studies were heterogeneously distributed, being more abundant for anchovy adults in the Western Mediterranean. Among all the environmental factors found significant, temperature, depth, salinity and chlorophyll-a concentration were the most studied for both species. Further emphasis was placed on the effect of these four variables, with various relationships found for the different life stages and areas. This review provides a general insight of what is known about how the environment influences growth, spawning, abundance and distribution of anchovy and sardine in the Mediterranean Sea, and identifies gaps of knowledge for future research. Our study sets a knowledge baseline for future studies about climate variability and change in the Mediterranean basin.

Introduction

European anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) are two of the most important commercial species in the Mediterranean Sea, reaching respectively up to 270,000 and 189,500 landed tons per year on average in the whole basin (FAO, 2018). These species are essential elements in the transfer of energy from lower to higher trophic level organisms (Palomera et al., 2007), coupling the Mediterranean pelagic and demersal environments. Cury et al. (2000) showed that fluctuations in small pelagic populations may modify ecosystem structure and functioning. Recent changes of anchovy and sardine have been observed in the Mediterranean Sea, in parallel with an overall increase of fishing effort (FAO, 2018).

Populations of anchovy and sardine are clearly influenced by changes in the environment (e.g. Lloret et al., 2004, Palomera et al., 2007). Environmental fluctuations can affect fish populations indirectly by controlling food quantity and quality or directly influencing annual recruitment, growth and condition of organisms (Basilone et al., 2017); with early life stages being particularly sensitive to these changes (Costalago et al., 2015). Sardine and anchovy have a relatively low position in the marine food web, and together with their short life-span and their reproductive strategy of producing large quantities of pelagic eggs, makes their populations strongly dependent on the environment (Palomera et al., 2007, Giannoulaki et al., 2013, Patti et al., 2020). This condition also makes them excellent bio-indicators of climate-driven changes in marine systems worldwide (Peck et al., 2013).

The study of the interactions of each of the two species with their environment is complex. Aspects such as interactions between both species, differences in habitat use, adaptations to the environmental change and the possible migratory movements in response to the climatic variability, pose an additional difficulty that needs to be considered and studied (Katara et al., 2011). The different studies available in the Mediterranean Sea on how environmental factors affect anchovy and sardine include different environmental variables and cover different ontogenetic stages and specific area. Therefore, the principal aim of this study was to systematically map and review information available on how environmental factors affect anchovy and sardine in the Mediterranean Sea in order to provide a general overview of what is known and identify the gaps of knowledge where future research should focus. The information extracted from the literature review is combined and discussed in terms of the relationships shown between the main environmental drivers at different life stages: egg, larvae, juveniles and adults. Since many marine resources from the Mediterranean Sea, including anchovy and sardine, are showing important declines (FAO, 2018), this review provides a step towards a better understanding on how marine species react to environmental factors, which is essential to manage marine resources sustainably under a context of climate change.

Section snippets

Materials and methods

We developed a bibliographic search of scientific articles, reviews and book chapters that studied the different environmental factors affecting sardine and anchovy populations in the Mediterranean Sea. A systematic review was applied following the PRISMA approach (Preferred Reporting Items for Systematic Reviews and Meta-Analyses; Moher et al., 2011), including all the information since 1990 until the cut-off date, 3rd of September 2020. This method includes three steps: (1) the systematic

Data distribution

A total of 1419 studies were finally screened of which 1416 records resulted by the systematic review, and 3 identified through other sources (i.e. grey literature) (Fig. 1). From the 160 articles assessed for eligibility, 113 were excluded because they did not analyse the relationship between the environmental factors and the anchovy/sardine parameters in a quantitative way and only 47 studies were selected to be included in the database (Table A.1; Appendix A). From those, 9 contained

Discussion

This paper has reviewed the effect of the main environmental factors found to be significantly related with anchovy or sardine variables: temperature, depth, salinity and chlorophyll.

Sea surface temperature appeared as one of the most important oceanographic variable influencing biological indicators, being a crucial factor affecting sardine and anchovy fluctuations (Katara et al., 2011). However, the effect of temperature changes with the ontogenetic stage and can even be contrasting (Basilone

Declaration of Competing Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Acknowledgements

This study was made under the auspices of PELWEB and SPELMED projects. PELWEB project is funded by the Spanish Ministry of Sciences and Innovation (ES-PN-2017-CTM2017-88939-R) and SPELMED project was funded by the EU DGMARE(SC NR 02-TENDER EASME/EMFF/2016/32) project. We specially thank Elena Lloret Lloret for her help with the maps.

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