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Mate fidelity and breeding site tenacity in a monogamous sandpiper, the black turnstone

We examined the relationship between mate fidelity and breeding site tenacity during a 5-year study of the black turnstone, Arenaria melanocephala, a socially monogamous sandpiper breeding in subarctic Alaska. We tested the predictions of several hypotheses regarding the incidence of divorce and the benefits of fidelity to mate and breeding site. Interannual return rates to the breeding grounds (88% for males, 79% for females) were among the highest yet recorded for any scolopacid sandpiper, and 88% of returning birds nested on their previous year's territory. The annual divorce rate was only 11%, and mate fidelity was significantly linked to fidelity to territory but independent of sex and year. Males arrived in spring significantly earlier than their mates and interannual fidelity was influenced by the relative timing of arrival of pair members. Reunited pairs had significantly higher fledging success than new pairs formed after death or divorce. The incidence of divorce was unrelated to reproductive success the previous year, although birds nested significantly further away after failure than after a successful nesting attempt. Sightings of marked individuals suggested that members of pairs do not winter together, and breeding site tenacity provides a mechanism through which pair members can reunite. We reject the 'incompatibility' hypothesis for divorce in turnstones, and our data contradict predictions of the 'better option' hypothesis. Alternatively, we propose the 'bet-hedging' hypothesis to explain the occurrence of divorce, which transpires when an individual pairs with a new mate to avoid the cost of waiting for a previous mate to return. Such costs can include remaining unmated, if the former mate has died, or experiencing lower reproductive success because of delayed breeding.

Animal Behaviour

Assessing conservation tools for an at-risk shorebird: Feasibility of headstarting for American Oystercatchers Haematopus palliatus

Management of threatened and endangered populations of wildlife increasingly relies upon active intervention such as predator control, habitat manipulation, and ex situ breeding or care. One tool that has received consideration for the management of declining or threatened avian populations is headstarting, or the artificial incubation of eggs and subsequent placement of newly hatched chicks in original or foster nests. We assessed the feasibility of implementing a headstarting program for the American Oystercatcher Haematopus palliatus , a species of high conservation concern in the eastern USA. Annual productivity is often low and lost during incubation, suggesting artificial incubation could enhance annual productivity. We used a control-impact approach to assign nests as either control or headstart and measured daily survival rate, success of parents accepting headstarted chicks, attendance patterns and behaviours of parents, and chick survival. We also implemented a transparent scoring process to rate the success of each step and the overall program. Daily survival rates of nests were significantly higher at headstart compared to control nests, and parents continued to incubate when eggs were well secured at nest sites. Attendance patterns and behaviour did not differ between headstart and control parents, and parents readily accepted healthy chicks whether they were returned to original or foster nests. Chick survival and subsequently annual productivity were, however, not higher at headstart compared to control nests suggesting that although we were able to enhance nest survival, low chick survival was still limiting annual productivity. Ultimately, headstarting may be most appropriate for American Oystercatchers where productivity is lost primarily to flooding, predation, or disturbance during the incubation stage but not during the chick-rearing stage. If, for example, high rates of nest loss are due to predators that also may prey upon chicks, then headstarting may not be an effective conservation tool.

Bird Conservation International

When a typical jumper skips: Itineraries and staging habitats used by Red Knots (Calidris canutus piersmai) migrating between northwest Australia and the New Siberian Islands

The ecological reasons for variation in avian migration, with some populations migrating across thousands of kilometres between breeding and non-breeding areas with one or few refuelling stops, in contrast to others that stop more often, remain to be pinned down. Red Knots Calidris canutus are a textbook example of a shorebird species that makes long migrations with only a few stops. Recognizing that such behaviours are not necessarily species-specific but determined by ecological context, we here provide a description of the migrations of a relatively recently described subspecies ( piersmai ). Based on data from tagging of Red Knots on the terminal non-breeding grounds in northwest Australia with 4.5- and 2.5-g solar-powered Platform Terminal Transmitters (PTTs) and 1.0-g geolocators, we obtained information on 19 route-records of 17 individuals, resulting in seven complete return migrations. We confirm published evidence that Red Knots of the piersmai subspecies migrate from NW Australia and breed on the New Siberian Islands in the Russian Arctic and that they stage along the coasts of southeastern Asia, especially in the northern Yellow Sea in China. Red Knots arrived on the tundra breeding grounds from 8 June onwards. Southward departures mainly occurred in the last week of July and the first week of August. We documented six non-stop flights of over c . 5000 km (with a maximum of 6500 km, lasting 6.6 days). Nevertheless, rather than staging at a single location for multiple weeks halfway during migration, piersmai -knots made several stops of up to a week. This was especially evident during northward migration, when birds often stopped along the way in southeast Asia and ‘hugged’ the coast of China, thus flying an additional 1000–1500 km compared with the shortest possible (great circle route) flights between NW Australia and the Yellow Sea. The birds staged longest in areas in northern China, along the shores of Bohai Bay and upper Liaodong Bay, where the bivalve Potamocorbula laevis , known as a particularly suitable food for Red Knots, was present. The use of multiple food-rich stopping sites during northward migration by piersmai is atypical among subspecies of Red Knots. Although piersmai apparently has the benefit of multiple suitable stopping areas along the flyway, it is a subspecies in decline and their mortality away from the NW Australian non-breeding grounds has been elevated.

New Siberian Islands

Why do some, but not all, tropical birds migrate? A comparative study of diet breadth and fruit preference

Annual migrations of birds profoundly influence terrestrial communities. However, few empirical studies examine why birds migrate, in part due to the difficulty of testing causal hypotheses in long-distance migration systems. Short-distance altitudinal migrations provide relatively tractable systems in which to test explanations for migration. Many past studies explain tropical altitudinal migration as a response to spatial and temporal variation in fruit availability. Yet this hypothesis fails to explain why some coexisting, closely-related frugivorous birds remain resident year-round. We take a mechanistic approach by proposing and evaluating two hypotheses (one based on competitive exclusion and the other based on differences in dietary specialization) to explain why some, but not all, tropical frugivores migrate. We tested predictions of these hypotheses by comparing diets, fruit preferences, and the relationships between diet and preference in closely-related pairs of migrant and resident species. Fecal samples and experimental choice trials revealed that sympatric migrants and residents differed in both their diets and fruit preferences. Migrants consumed a greater diversity of fruits and fewer arthropods than did their resident counterparts. Migrants also tended to have slightly stronger fruit preferences than residents. Most critically, diets of migrants more closely matched their preferences than did the diets of residents. These results suggest that migrants may be competitively superior foragers for fruit compared to residents (rather than vice versa), implying that current competitive interactions are unlikely to explain variation in migratory behavior among coexisting frugivores. We found some support for the dietary specialization hypothesis, propose refinements to the mechanism underlying this hypothesis, and discuss how dietary specialization might ultimately reflect past interspecific competition. We recommend that future studies quantify variation in nutritional content of tropical fruits, and determine whether frugivory is a consequence or a cause of migratory behaviour.

Evolutionary Ecology

Non-random pairing in American kestrels: mate choice versus intra-sexual competition

Natural selection may influence the arrangement of individuals into mated pairs through either inter-sexual (mate choice) or intra-sexual selection (competition). A study of the American kestrel, Falco sparverius , in northern Saskatchewan distinguished between these two processes using size as a measure of the bird's competitive ability, and condition (mass scaled to body size) as an index of quality. Both sexes arrive on the study area after spring migration in equal numbers and males establish territories. Males and females that moved among territories at the time of pair formation were not different in size or condition from those that did not move, suggesting that birds were not being displaced by superior competitors, and that females moved to encounter potential mates. Within mated pairs, there was no relationship between a bird's size and the condition of its mate for either sex as would be predicted if intra-sexual competitition explained mating patterns. Instead, there was positive assortative mating by condition, suggesting that both sexes used quality as the criterion in choosing mates. There was no correlation between the sizes of males and females in mated paird. Because there were no differences in size or condition of breeding and non-breeding males, factors other than physical attributes, such as prior experience with the area, may determine a male's success in obtaining a territory. Because females that did not obtain mates were in poorer condition than those that did, males may have rejected poor quality females. The results suggest that intra-sexual competition was not important for pair formation, and that kestrels chose mates on the basis of quality.

Animal Behaviour

Migratory hummingbirds make their own rules: The decision to resume migration along a barrier

Knowing how naïve migrants respond to intrinsic and extrinsic factors experienced en route will allow a more thorough understanding of the endogenous migratory programme. To understand how inexperienced individuals respond to ecological features, we tracked the migratory departures of juvenile ruby-throated hummingbirds, Archilochus colubris , one of the smallest (∼ 3 g) and least-studied migrants, along the Gulf of Mexico during southbound migration using an international automated radiotelemetry system. The recent miniaturization of radiotags provides a novel method to track one of the smallest migratory birds, rendering the first information on departure decisions of known hummingbirds in relation to an ecological barrier. Using weather conditions and individual attributes, we also determined which factors influenced the time and direction of departure from a coastal stopover site. Most migrants (83%) departed in the morning, and daily departure time was only influenced by stopover duration, the amount of time spent at a stopover site. The majority (77%) of departure orientations paralleled the coastline, and we found little influence of any factor on departure direction. Our results reveal that (1) juvenile hummingbirds departing coastal Alabama move in a direction indicative of a circum-Gulf path during southbound migration and (2) departure decisions support a fly-and-forage strategy in which hummingbirds likely take advantage of resources along the coast while moving towards their destination.

Gulf of Mexico

Birds choose long-term partners years before breeding

Pair bonds can provide social benefits to long-term monogamous species alongside their benefits for reproduction. However, little is known about when these bonds form, in particular how long they are present before breeding. Previous studies of pair formation in long-term monogamous birds have been rather data-limited, but for many migratory birds they report pair formation on the wintering grounds. We provide the first systematic investigation of prebreeding association patterns of long-term monogamous pairs by examining entire life histories based on tracking data of migratory whooping cranes, Grus americana. We found that a substantial portion (62%) of breeding pairs started associating at least 12 months before first breeding, with 16 of 58 breeding pairs beginning to associate over 2 years before first breeding. For most pairs, these associations with future breeding partners also became unique and distinguishable from association patterns with nonpartner individuals 12 months before first breeding. In addition, 60% of pair associations began before at least one partner had reached nominal sexual maturity. Most pairs began associating in the late spring upon arrival at the summer grounds, while associations beginning at other times of the year were rare. Patterns in the associations of pairs prior to breeding can point to the potential benefits of prebreeding relationships, for instance providing support in competitive interactions or increasing partner familiarity.

Animal Behaviour

Forced copulation results in few extrapair fertilizations in Ross's and lesser snow geese

Extrapair paternity varies from 0 to over 70% of young among various populations of birds. Comparative studies have suggested that this variation is related to nesting density, breeding synchrony and the proportion of extrapair copulations. We used minisatellite DNA fingerprinting to examine levels of extrapair paternity in Ross's geese, Chen rossi, and lesser snow geese, C. caerulescens c. (hereafter snow geese) nesting in the largest known goose colony in the world. These geese have one of the highest known percentages of extrapair copulation (46-56% of all attempted copulations), and all of these appeared to be forced. Among all successful copulations, 33 and 38% were extrapair in Ross's and snow geese, respectively. Despite the high percentage of extrapair copulations, extrapair paternity was low in both Ross's and snow geese (2-5% of young). Extrapair paternity was not related to nest density in either species. However, in snow geese, extrapair paternity was more likely to occur in nests of females that nested asynchronously, either early or late in the season. This is one of a few reported examples of a negative relationship between extrapair paternity and breeding synchrony. Extrapair young also tended to come from eggs laid later in the clutch. Although forced extrapair copulations appear to be a relatively inefficient reproductive tactic for males, they may provide a reproductive advantage for some males.

Animal Behaviour

Certainty of paternity and paternal investment in eastern bluebirds and tree swallows

Extra-pair paternity is common in many socially monogamous passerine birds with biparental care. Thus, males often invest in offspring to which they are not related. Models of optimal parental investment predict that, under certain assumptions, males should lower their investment in response to reduced certainty of paternity. We attempted to reduce certainty of paternity experimentally in two species, the eastern bluebird, S ialia sialis , and the tree swallow, Tachycineta bicolor , by temporarily removing fertile females on two mornings during egg laying. In both species, experimental males usually attempted to copulate with the female immediately after her reappearance, suggesting that they experienced the absence of their mate as a threat to their paternity. Experimental males copulated at a significantly higher rate than control males. However, contrary to the prediction of the model, experimental males did not invest less than control males in their offspring. There was no difference between experimental and control nests in the proportion of male feeds, male and female feeding rates, nestling growth and nestling condition and size at age 14 days. We argue that females might have restored the males’ confidence in paternity after the experiment by soliciting or accepting copulations. Alternatively, males may not reduce their effort, because the fitness costs to their own offspring may outweigh the benefits for the males, at least in populations where females cannot fully compensate for reduced male investment.

Animal Behaviour