Abstract
The Gulf of Lions has faced a sharp drop in the catches of its two main small pelagic exploited species, the sardine Sardina pilchardus and the anchovy Engraulis encrasicolus since the mid-2000s, despite both population abundances remaining high. This situation has been due to a severe decrease in individual body condition and size as a result of both lower growth and the disappearance of the oldest and largest individuals. While overfishing, predation or disease outbreaks have been refuted to explain this situation, one major hypothesis remained to be investigated. A potential shift in sardine and anchovy diet towards smaller planktonic prey indeed suggested bottom-up control as the main driver of these populations in the Gulf of Lions. The first aim of this thesis was to investigate whether bottom-up processes could explain the changes in sardine growth and condition through changes in both food size and/or quantity and to understand the behavioral and physiological mechanisms involved in this control. The second objective of this PhD thesis was to identify the potential underlying drivers leading to adult overmortality. To do so, we combined an experimental approach on wild sardines maintained in captivity with a modeling approach. Experimentations showed that body condition, growth and storage lipids were significantly impacted by both food size and quantity. Thus, sardines fed on small particles needed to consume twice as much as those feeding on large particles to achieve the same condition and growth. Such results seemed to be linked to higher energy expenditures of sardines while filtering small prey compared to particulate feeding on large prey (sardines being able to shift between two feeding modes according to the prey size). Moreover, our results suggested several adaptations to cope with small food and caloric restriction. The study of the gill raker apparatus involved in the filtration of small prey suggested an increase of the filtration capacity for a given length between 2007-2009 and 2016. Then, sardines fed on small particles exhibited higher mitochondria efficiency and abundance suggesting energy-saving adaptation. Finally, sardines accustomed to feed on small pellets showed lower activity to limit energy expenditure. Nevertheless, all these strategies might incur other costs or may not be enough to compensate the high energy demands of filtration on small prey, as growth and condition remained lower for sardines filtering small prey in all our experiments. Further, sardines fed on large pellets exhibited higher spawning frequency than sardines fed with the same quantity of small ones. The low egg production of these sardines might be explained by a too high body condition of these individuals to observe a change in energy trade-off towards reproduction. For the same reasons, small particle meals did not seem to impact their immunity and stress, leucocyte and cortisol concentrations being similar whatever the feeding treatment. Furthermore, to investigate the hypothesis of adult overmortality, we first studied whether individual could die from starvation and low body reserves. The survival probability sharply decreased when the body condition index became lower than 0.75 and the threshold of 0.72 was identified as the entry in phase III of fasting. While the proportion of sardines reaching such thresholds in the wild remains low, it still increased two-fold in the recent period, reaching about 10% in winter months. A DEB model parameterized using a combination of in-situ and experimental data suggested a lower survival probability for larger fish. Individuals larger than 14 cm, i.e. older than 2-3 years, had a lower than 50 % probability to survive 1 month after the reproduction period. In conclusion, these previous results comforted the two hypotheses of a bottom-up control and an overmortality of adult sardines after reproduction to explain the dynamic and demographic truncation of the sardine population.