No abstract

Migratory birds need considerable energy reserves to fuel long-distance flights to their breeding grounds in spring. To attain sufficient energy deposits before departure, birds require high daily intake rates, which can be reached by utilizing high-quality food. During such periods of high energy demand, animals often track changes in the nutritious value of their food, for example by switching to a more profitable habitat or diet. Pre-migratory Barnacle Geese Branta leucopsis staging along the Wadden Sea coast are known to switch from pastures to salt marshes during spring. Previous studies have suggested that Barnacle Geese switch habitat to track changes in protein levels, which drop in pastures as spring progresses, and to avoid high levels of disturbance. Here we made use of detailed information on pre-migratory habitat use of individual Barnacle Geese tracked by GPS/accelerometer tags to assess which factors may drive a habitat switch. We analysed habitat use and time budgets of individual birds and combined this with data on food quality in two habitats to analyse differences in food intake. We found large individual variation in pre-migratory habitat use, both in the extent of salt marsh use, as well as in the timing of a switch from pastures to salt marshes. In salt marshes, geese spent more time grazing and made fewer flight movements, potentially as they experienced lower levels of disturbance compared to geese in pastures. By increasing grazing time and reducing flight movements, geese in salt marshes may compensate for reduced food quality. Our results show that Barnacle Geese trade-off high intake rates and high costs in pastures with low intake rates and low costs in salt marshes.

Capsule: The number of breeding pairs of Europe's largest Arctic Skua Stercorarius parasiticus colony at Slettnes, Norway, showed a dramatic decline of at least 50% over two decades, with food shortage in four years and increasing predation by Red Fox Vulpes vulpes leading to total breeding failure in five out of six recent study years. Aims: To document the decline of Europe's largest Arctic Skua colony and quantify bottom-up and top-down effects on reproduction. Methods: We compared nest counts between 1997-1998 and 2014-2019 and collected data on egg size, clutch size and nest success for all years, and adult body mass, nest attendance, at-sea activity, aggressive nest defence, Red Fox Vulpes vulpes encounters, daily nest survival and adult survival for 2014-2019. We deployed nest cameras to identify predators in 2018-2019. In addition, we developed a demographic model to estimate the fecundity required for a stable population. Results: A higher proportion of time spent at sea, small eggs, low adult female body mass and indirect assessment of foraging fish availability suggested food shortages in four of six recent study years. At the same time, nest predation by Red Foxes, the likely predator involved, increased during the six-year study. The combined effects of food shortage and nest predation led to total breeding failures in 2017-2019. Conclusion:We provide evidence of both bottom-up (food shortage) and top-down (predation) effects on reproductive investment and hatching success in this colony. The reproductive output in recent years is far too low to sustain a stable population. The severe decline of the Arctic Skua colony at Slettnes fits reported trends for this species across most of its European breeding range, as well as for its important host species, the Arctic Tern Sterna paradisaea and the Black-legged Kittiwake Rissa tridactyla.

Females should facultatively bias offspring sex ratio when fitness returns vary among sexes. In cooperative breeders, where individuals help raise others’ young, overproducing the philopatric sex will be adaptive when helpers are absent, whereas overproducing the dispersive sex may be adaptive to reduce intrasexual competition. Thus, fitness returns are expected to vary with the social environment. However, any offspring sex-ratio biases may also result from consistent among-female differences (e.g. quality) and/or environmental variables (e.g. food availability). Yet, few studies have disentangled facultative from persistent biases. We investigated offspring sex-ratio biases in relation to the social environment in cooperatively breeding red-winged fairy-wrens (Malurus elegans). Repeated observations of the same females over nine years allowed for disentanglement of facultative from persistent biases. Females without help did not overproduce daughters, despite female helpers being associated with higher fledgling survival (resource enhancement hypothesis). Instead, females without helpers facultatively overproduced sons —the slower dispersing sex— thereby ensuring long-term helper availability. Furthermore, offspring sex ratio was not biased towards the rarer sex of helpers present in the group or population (resource competition hypothesis). However, females with sex-biased helping produced similarly skewed offspring sex ratios. This among-female association may not be surprising, because helpers are previous seasons’ offspring. Thus, in addition to facultative responses to prevailing social conditions, we found evidence for persistent biases among females. This could potentially explain previous evidence for resource competition/enhancement that have typically been interpreted as facultative responses, highlighting the need for a within-female approach to better understand the adaptiveness of sex-ratio biases. Significance statement Under certain conditions, females may benefit from producing a biased offspring sex ratio, but evidence for such effects in vertebrates is weak and inconsistent. Here, using observations of the same females under different social conditions, we show that cooperatively breeding red-winged fairy-wrens facultatively biased offspring sex ratio towards sons when living in pairs, thereby ensuring the availability of a workforce to assist in raising future offspring. However, biased offspring sex ratio patterns may also be the result of consistent differences among females. Indeed, we also found evidence for such patterns and suggest that this could be an explanation for previous findings which are often interpreted as facultative responses.