Spawner weight and ocean temperature drive Allee effect <?xmltex \hack{\break}?> dynamics in Atlantic cod, Gadus morhua: <?xmltex \hack{\break}?> inherent and emergent density regulation

Stocks of Atlantic cod, Gadus morhua, show diverse recovery responses when fishing pressure is relieved. The expected outcome of reduced fishing pressure is that the population regains its size. However, there are also cod stocks that seem to be locked in a state of low abundance from which population growth does not occur (or only slowly occurs). A plausible explanation for this phenomenon can be provided by the Allee effect, which takes place when recruitment per capita is positively related to population density or abundance. However, because of methodological limitations and data constraints, such a phenomenon is often perceived as being rare or non-existent in marine fish. In this study, we used time series of 17 Atlantic cod stocks to fit a family of population equations that consider the abundance of spawners, their body weight and sea water temperature as independent components of recruitment. The developed stock-recruitment function disentangles the effects of spawner abundance, spawner weight and temperature on recruitment dynamics and captures the diversity of density dependencies (compensation, Allee effect) of the recruitment production in Atlantic cod. The results show for 13 cod stocks an inherent spawner-abundance-related Allee effect. Allee effect strength, i.e., the relative change between maximum and minimum recruitment per capita at low abundance, was increased when recruitment production was suppressed by unfavorable changes in water temperature and/or in spawner weight. The latter can be a concomitant of heavy fishing or a result of temperature-related altered body growth. Allee effect strength was decreased when spawner weight and/or temperature elevated recruitment production. We show how anthropogenic stress can increase the risk of Allee effects in stocks where ocean temperature and/or spawner weight had been beneficial in the past but are likely to unmask and strengthen an inherent Allee effect under future conditions.