Evidence that tufted puffins Fratercula cirrhata use colony overflights to reduce kleptoparasitism risk

Predation, foraging and mating costs are critical factors shaping life histories. Among colonial seabirds, colony overflights may enhance foraging or mating success, or diminish the risk of predation and kleptoparasitism. The latter possibility is difficult to test because low predation or kleptoparasitism rates could be due either to low danger or to effective counter-tactics by prey. Tufted puffins Fratercula cirrhata breeding at a large colony in British Columbia, Canada, deliver several loads of fish each day to their nestlings and are targets for kleptoparasitism by glaucous-winged gulls Larus glaucescens . In the present study, we documented the ecological conditions under which overflights occurred in order to assess when overflights were made and to statistically isolate the effect of overflights on kleptoparasitism risk at this site. Load-carrying puffins engaged in overflights under ecological conditions associated with relatively high rates of kleptoparasitism. Further, when ecological factors determining risk were statistically controlled, overflights were correlated with marginally lower chances of kleptoparasitism than when the risk factors were ignored. The results suggest that breeding puffins at this site use overflights for kleptoparasite avoidance. This tactic is used sparingly, suggesting it is costly. Costs of overflight behaviour might contribute to the impact of kleptoparasitism on the breeding success of tufted puffins.     

Protandrous arrival timing to breeding areas: a review

Protandry, the earlier arrival of males to breeding areas than females, is a common pattern of sex-biased timing in many animal taxa (e.g. some insects, fish, amphibians, reptiles, birds and mammals). The adaptive significance of protandry is not fully understood and, since the 1970s, at least seven hypotheses for protandry have been proposed. We describe each of these hypotheses and summarize what is known about each. In three of these hypotheses, the relative arrival timing of males and females has no direct fitness consequences for males or females, but selection for different timing in each sex indirectly produces protandry. In the other four hypotheses, the difference between male and female timing has fitness consequences for males or females and selection directly maintains the fitness-maximizing degree of sex-biased timing. The hypotheses are not mutually exclusive, and the degree of multiple mating by males and the occurrence of male territoriality seem to determine the relative importance of each hypothesis. In order to understand the adaptive significance of sex-biased timing, future studies need to consider all the alternatives and to assess the costs and benefits to males of early arrival relative to calendar date, to other males and to females.     

Range‐wide patterns of migratory connectivity in the western sandpiper Calidris mauri

Understanding the population dynamics of migratory animals and predicting the consequences of environmental change requires knowing how populations are spatially connected between different periods of the annual cycle. We used stable isotopes to examine patterns of migratory connectivity across the range of the western sandpiper Calidris mauri. First, we developed a winter isotope basemap from stable‐hydrogen (δD), ‐carbon (δ¹³C), and ‐nitrogen (δ¹⁵N) isotopes of feathers grown in wintering areas. δD and δ¹⁵N values from wintering individuals varied with the latitude and longitude of capture location, while δ¹³C varied with longitude only. We then tested the ability of the basemap to assign known‐origin individuals. Sixty percent of wintering individuals were correctly assigned to their region of origin out of seven possible regions. Finally, we estimated the winter origins of breeding and migrant individuals and compared the resulting empirical distribution against the distribution that would be expected based on patterns of winter relative abundance. For breeding birds, the distribution of winter origins differed from expected only among males in the Yukon‐Kuskokwim (Y‐K) Delta and Nome, Alaska. Males in the Y‐K Delta originated overwhelmingly from western Mexico, while in Nome, there were fewer males from western North America and more from the Baja Peninsula than expected. An unexpectedly high proportion of migrants captured at a stopover site in the interior United States originated from eastern and southern wintering areas, while none originated from western North America. In general, we document substantial mixing between the breeding and wintering populations of both sexes, which will buffer the global population of western sandpipers from the effects of local habitat loss on both breeding and wintering grounds.     

Effects of predation danger on migration strategies of sandpipers

We examine the potential selective importance of predation danger on the evolution of migration strategies of arctic‐breeding calidrid sandpipers. Adult calidrids truncate parental care for reasons not obviously related to levels of food abundance on the breeding areas or at migratory stopover sites, suggesting that a different trade‐off occurs between providing additional care and adult survivorship. The southward migrations of adult western sandpipers precede those of migratory peregrine falcons by almost a month. By moving early and quickly, adults remain ahead of migrant falcons all the way to their non‐breeding areas, where they rapidly moult flight feathers. They complete the moult just as falcons arrive in late September–October. By migrating early, they avoid exposure to falcons when they are unusually vulnerable, due to the requirements for fuelling migratory flight and of wing feather moult. Juvenile western sandpipers migrate south just as falcon numbers start to increase, but do not moult flight feathers in their first winter. Pacific dunlin use an alternative strategy of remaining and moulting in Alaska after falcons depart, and migrating to their overwintering sites after migrants have passed. East of the Rocky Mountains, the southbound migration of falcons begins 4–6 weeks later. Southbound semipalmated sandpipers make extended migratory stopovers, but their lengths of stay shorten prior to falcon migration to the sites in September. Predation danger also may affect the evolution of migration routes. Southbound western sandpipers fly directly from Alaska to southern British Columbia, in contrast to the multi‐stage journey northward along the Alaska panhandle. We estimate that a direct flight would be more economical on northward migration, but may be avoided because it would expose sandpipers to higher mass‐dependent predation danger from migratory falcons, which travel north with sandpipers. By contrast, few raptors are present in Alaska during preparation for the southward flight. A temporal and spatial window of safety may also permit semipalmated sandpipers to become extremely vulnerable while preparing for trans‐Atlantic southward flights. Danger management may account for the these previously enigmatic features of calidrid migration strategies, and aspects of those of other birds.     

Western sandpipers have altered migration tactics as peregrine falcon populations have recovered

The presence of top predators can affect prey behaviour, morphology and life history, and thereby can produce indirect population consequences greater and further reaching than direct depredation would have alone. Raptor species in the Americas are recovering since restrictions on the use of dichlorodiphenyltrichloroethane (DDT) and the implementation of conservation measures, in effect constituting a hemisphere-wide predator-reintroduction experiment, and profound effects on populations of their prey are to be expected. Here, we document changes in the behaviour of western sandpipers (Calidris mauri) at migratory stopover sites over two decades. Since 1985, migratory body mass and stopover durations of western sandpipers have fallen steadily at some stopovers in the Strait of Georgia, British Columbia. Comparisons between years, sites and seasons strongly implicate increasing danger from the recovery of peregrine falcons (Falco peregrinus) as a causal factor. A decade-long ongoing steep decline in sandpiper numbers censused on our study site is explained entirely by the shortening stopover duration, rather than fewer individuals using the site. Such behavioural changes are probably general among migratory shorebird species, and may be contributing to the widespread census declines reported in North America.     

Rapid adaptive adjustment of parental care coincident with altered migratory behaviour

The optimal duration of parental care is shaped by the trade-off between investment in current and expected future reproductive success. A change in migratory behaviour is expected to affect the optimal duration of parental care, because migration and non-migration differ in expectations of future reproductive success as a result of differential adult and/or offspring mortality. Here we studied how a recent emergence of non-migratory behaviour has affected the duration of parental care in the previously (until the 1980s) strictly migratory Russian breeding population of the barnacle geese Branta leucopsis. As a measure of parental care, we compared the vigilance behaviour of parents and non-parents in both migratory and non-migratory barnacle geese throughout the season. We estimated the duration of parental care at 233 days for migratory and 183 days for non-migratory barnacle geese. This constitutes a shortening of the duration of parental care of 21% in 25 years. Barnacle geese are thus able to rapidly adapt their parental care behaviour to ecological conditions associated with altered migratory behaviour. Our study demonstrates that a termination of migratory behaviour resulted in a drastic reduction in parental care and highlights the importance of studying the ecological and behavioural consequences of changes in migratory behaviour and the consequences of these changes for life-history evolution.