Strong survival selection on seasonal migration versus residence induced by extreme climatic events.

Elucidating the full eco-evolutionary consequences of climate change requires quantifying the impact of extreme climatic events (ECEs) on selective landscapes of key phenotypic traits that mediate responses to changing environments. Episodes of strong ECE-induced selection could directly alter population composition, and potentially drive micro-evolution. However, to date, few studies have quantified ECE-induced selection on key traits, meaning that immediate and longer-term eco-evolutionary implications cannot yet be considered. One widely expressed trait that allows individuals to respond to changing seasonal environments, and directly shapes spatio-seasonal population dynamics, is seasonal migration versus residence. Many populations show considerable among-individual phenotypic variation, resulting in 'partial migration'. However, variation in the magnitude of direct survival selection on migration versus residence has not been rigorously quantified, and empirical evidence of whether seasonal ECEs induce, intensify, weaken or reverse such selection is lacking. We designed full annual cycle multi-state capture-recapture models that allow estimation of seasonal survival probabilities of migrants and residents from spatio-temporally heterogeneous individual resightings. We fitted these models to 9 years of geographically extensive year-round resighting data from partially migratory European shags Phalacrocorax aristotelis. We thereby quantified seasonal and annual survival selection on migration versus residence across benign and historically extreme non-breeding season (winter) conditions, and tested whether selection differed between females and males. We show that two of four observed ECEs, defined as severe winter storms causing overall low survival, were associated with very strong seasonal survival selection against residence. These episodes dwarfed the weak selection or neutrality evident otherwise, and hence caused selection through overall annual survival. The ECE that caused highest overall mortality and strongest selection also caused sex-biased mortality, but there was little overall evidence of sex-biased selection on migration versus residence. Our results imply that seasonal ECEs and associated mortality can substantially shape the landscape of survival selection on migration versus residence. Such ECE-induced phenotypic selection will directly alter migrant and resident frequencies, and thereby alter immediate spatio-seasonal population dynamics. Given underlying additive genetic variation, such ECEs could potentially cause micro-evolutionary changes in seasonal migration, and thereby cause complex eco-evolutionary population responses to changing seasonal environments.

What, where and when: deconstructing memory.

The ability of animals to remember the what, where and when of a unique past event is used as an animal equivalent to human episodic memory. We currently view episodic memory as reconstructive, with an event being remembered in the context in which it took place. Importantly, this means that the components of a what, where, when memory task should be dissociable (e.g. what would be remembered to a different degree than when). We tested this hypothesis by training hummingbirds to a memory task, where the location of a reward was specified according to colour (what), location (where), and order and time of day (when). Although hummingbirds remembered these three pieces of information together more often than expected, there was a hierarchy as to how they were remembered. When seemed to be the hardest to remember, while errors relating to what were more easily corrected. Furthermore, when appears to have been encoded as a combination of time of day and sequence information. As hummingbirds solved this task using reconstruction of different memory components (what, where and when), we suggest that similar deconstructive approaches may offer a useful way to compare episodic and episodic-like memories.