Where east meets west: Phylogeography of the high Arctic North American brant goose.

Genetic variation in Arctic species is often influenced by vicariance during the Pleistocene, as ice sheets fragmented the landscape and displaced populations to low- and high-latitude refugia. The formation of secondary contact or suture zones during periods of ice sheet retraction has important consequences on genetic diversity by facilitating genetic connectivity between formerly isolated populations. Brant geese (Branta bernicla) are a maritime migratory waterfowl (Anseriformes) species that almost exclusively uses coastal habitats. Within North America, brant geese are characterized by two phenotypically distinct subspecies that utilize disjunct breeding and wintering areas in the northern Pacific and Atlantic. In the Western High Arctic of Canada, brant geese consist of individuals with an intermediate phenotype that are rarely observed nesting outside this region. We examined the genetic structure of brant geese populations from each subspecies and areas consisting of intermediate phenotypes using mitochondrial DNA (mtDNA) control region sequence data and microsatellite loci. We found a strong east-west partition in both marker types consistent with refugial populations. Within subspecies, structure was also observed at mtDNA while microsatellite data suggested the presence of only two distinct genetic clusters. The Western High Arctic (WHA) appears to be a secondary contact zone for both Atlantic and Pacific lineages as mtDNA and nuclear genotypes were assigned to both subspecies, and admixed individuals were observed in this region. The mtDNA sequence data outside WHA suggests no or very restricted intermixing between Atlantic and Pacific wintering populations which is consistent with published banding and telemetry data. Our study indicates that, although brant geese in the WHA are not a genetically distinct lineage, this region may act as a reservoir of genetic diversity and may be an area of high conservation value given the potential of low reproductive output in this species.

Environmental conditions during breeding modify the strength of mass-dependent carry-over effects in a migratory bird.

In many animals, processes occurring in one season carry over to influence reproductive success and survival in future seasons. The strength of such carry-over effects is unlikely to be uniform across years, yet our understanding of the processes that are capable of modifying their strength remains limited. Here we show that female light-bellied Brent geese with higher body mass prior to spring migration successfully reared more offspring during breeding, but only in years where environmental conditions during breeding were favourable. In years of bad weather during breeding, all birds suffered reduced reproductive output irrespective of pre-migration mass. Our results suggest that the magnitude of reproductive benefits gained by maximising body stores to fuel breeding fluctuates markedly among years in concert with conditions during the breeding season, as does the degree to which carry-over effects are capable of driving variance in reproductive success among individuals. Therefore while carry-over effects have considerable power to drive fitness asymmetries among individuals, our ability to interpret these effects in terms of their implications for population dynamics is dependent on knowledge of fitness determinants occurring in subsequent seasons. 

Heterozygosity-fitness correlations in a migratory bird: an analysis of inbreeding and single-locus effects.

Studies in a multitude of taxa have described a correlation between heterozygosity and fitness and usually conclude that this is evidence for inbreeding depression. Here, we have used multilocus heterozygosity (MLH) estimates from 15 microsatellite markers to show evidence of heterozygosity-fitness correlations (HFCs) in a long-distance migratory bird, the light-bellied Brent goose. We found significant, positive heterozygosity-heterozygosity correlations between random subsets of the markers we employed, and no evidence that a model containing all loci as individual predictors in a multiple regression explained significantly more variation than a model with MLH as a single predictor. Collectively, these results lend support to the hypothesis that the HFCs we have observed are a function of inbreeding depression. However, we do find that fitness correlations are only detectable in years where population-level productivity is high enough for the reproductive asymmetry between high and low heterozygosity individuals to become apparent. We suggest that lack of evidence of heterozygosity-fitness correlations in animal systems may be because heterozygosity is a poor proxy measure of inbreeding, especially when employing low numbers of markers, but alternatively because the asymmetries between individuals of different heterozygosities may only be apparent when environmental effects on fitness are less pronounced.

Cultural inheritance drives site fidelity and migratory connectivity in a long-distance migrant.

Cultural transmission is thought to be a mechanism by which migratory animals settle into habitats, but little evidence exists in wild populations because of the difficulty of following individuals over successive generations and wide geographical distances. Cultural inheritance of migration routes represents a mechanism whereby geographical isolation can arise between separate groups and could constrain individuals to potentially suboptimal sites within their range. Conversely, adopting the parental migratory route in adult life, rather than dispersing randomly, may increase an individual's reproductive success because that strategy has already been proven to allow successful breeding. We combined a pedigree of related light-bellied Brent geese (Branta bernicla hrota) with 6 years of observations of marked birds to calculate the dispersal distances of adult offspring from their parents in both Ireland and Iceland. In both countries, the majority of offspring were found to recruit into or near their parental sites, indicating migratory connectivity in the flyway. Despite this kin structure, we found no evidence of genetic differentiation using genotype data from 1127 individuals across 15 microsatellite loci. We suggest that the existence of migratory connectivity of subpopulations is far more common than previous research indicates and that cultural information may play an important role in structuring reproductive isolation among them.

Carry-over effects reveal reproductive costs in a long-distance migrant.

1. It has been known for some time that the consequences of 'decisions' made at one point in an animal's life may not always be borne immediately. For example, numerous studies have demonstrated the trade-off between current and future breeding success across multiple taxa. 2. It is becoming increasingly clear that such processes may also operate among seasons, such that the conditions experienced at one point in the annual cycle may have significant downstream impacts, or 'carry-over effects', and this is particularly evident among migratory species. We might therefore predict that certain combinations of reproductive and migratory strategy could lead to profound carry-over effects. However, the extent to which these phenomena might generate variation in fitness within a population is unclear. 3. Here, we investigate how winter habitat selection in a long-distance migrant, with extended parental care (the Light-bellied Brent goose) is influenced by parental status and how this has a counterintuitive effect on subsequent breeding success. 4. Dominant individuals and groups generally monopolize the best quality resources. In the case of geese, families are dominant; however, our findings highlight a hidden cost to raising a family. Stable isotope analysis demonstrates that later in the non-breeding season, adults with families utilize lower quality resources than non-breeders. This is probably caused by parents being constrained in habitat choice by the lower foraging efficiency of their juveniles. Consequently, parental adults end the winter in poorer condition than non-breeders. 5. We further demonstrate that parents in one year are less likely than expected to breed again in the next year and suggest that this is caused by conditions during the non-breeding period being carried over into the breeding season. In conclusion, we demonstrate previously hidden costs to raising a family, which are likely to be important in terms of life-history evolution.

Bioaccumulation and biomagnification of organochlorines in a marine food web at a pristine site in Iceland.

Organochlorine (OC) bioaccumulation and biomagnification were studied in a marine food web at a pristine site in Iceland. The species studied were the gastropod and grazer chink shell (Lacuna vincta), the filter feeding bivalve blue mussel (Mytilus edulis), the predators butterfish (Pholis gunnellus), and the seabird black guillemot (Cepphus grylle), all sampled and analysed in 1996-1997. Individual OC levels were generally low in chink shell and blue mussels, somewhat elevated in the fish, and an order of a magnitude higher in the top predator black guillemot, except for Sigma HCH (hexachlorocyclohexane isomers) and Sigma chlordane levels, which were similar in all organisms, ranging from 10 to 36 ng/g lipid weight (lw). In the molluscs and fish, mean concentrations of Sigma PCB (polychlorinated biphenyl) ranged from 111 to 377 ng/g lw, Sigma DDT (dichlorodiphenyltrichloroethane) ranged from 19 to 65 ng/g lw, and HCB (hexachlorobenzene) ranged from 21 to 30 ng/g lw. The levels of same OCs in the black guillemot were on average 2352, 361, and 283 ng/g lw, respectively. The OC tissue concentrations in blue mussel and black guillemot are comparable to levels in Arctic and sub-Arctic regions, but OC levels in blue mussel tissue were an order of magnitude lower than found in the North Sea and the Baltic Sea. The relative composition of OCs were generally similar among species with the PCB congeners emerging as the most abundant compounds with levels an order of magnitude higher than the other compounds in all species. Food web magnification factors (FWMFs) were determined for the OCs by using trophic levels determined from delta(15)N. FWMFs >1, indicating biomagnification, were found for Sigma PCB, penta- or higher chlorinated PCBs (e.g., PCB 101, 118, 138, 153, 180), beta-HCH, HCB, Sigma DDT, p,p-DDE, and transnonachlor. The highest FWMF was observed for PCB 180 at FWMF = 5.8.

A polar system of intercontinental bird migration.

Studies of bird migration in the Beringia region of Alaska and eastern Siberia are of special interest for revealing the importance of bird migration between Eurasia and North America, for evaluating orientation principles used by the birds at polar latitudes and for understanding the evolutionary implications of intercontinental migratory connectivity among birds as well as their parasites. We used tracking radar placed onboard the ice-breaker Oden to register bird migratory flights from 30 July to 19 August 2005 and we encountered extensive bird migration in the whole Beringia range from latitude 64 degrees N in Bering Strait up to latitude 75 degrees N far north of Wrangel Island, with eastward flights making up 79% of all track directions. The results from Beringia were used in combination with radar studies from the Arctic Ocean north of Siberia and in the Beaufort Sea to make a reconstruction of a major Siberian-American bird migration system in a wide Arctic sector between longitudes 110 degrees E and 130 degrees W, spanning one-third of the entire circumpolar circle. This system was estimated to involve more than 2 million birds, mainly shorebirds, terns and skuas, flying across the Arctic Ocean at mean altitudes exceeding 1 km (maximum altitudes 3-5 km). Great circle orientation provided a significantly better fit with observed flight directions at 20 different sites and areas than constant geographical compass orientation. The long flights over the sea spanned 40-80 degrees of longitude, corresponding to distances and durations of 1400-2600 km and 26-48 hours, respectively. The birds continued from this eastward migration system over the Arctic Ocean into several different flyway systems at the American continents and the Pacific Ocean. Minimization of distances between tundra breeding sectors and northerly stopover sites, in combination with the Beringia glacial refugium and colonization history, seemed to be important for the evolution of this major polar bird migration system.

Population-scale drivers of individual arrival times in migratory birds.

1. In migratory species, early arrival on the breeding grounds can often enhance breeding success. Timing of spring migration is therefore a key process that is likely to be influenced both by factors specific to individuals, such as the quality of winter and breeding locations and the distance between them, and by annual variation in weather conditions before and during migration. 2. The Icelandic black-tailed godwit Limosa limosa islandica population is currently increasing and, throughout Iceland, is expanding into poorer quality breeding areas. Using a unique data set of arrival times in Iceland in different years for individuals of known breeding and wintering locations, we show that individuals breeding in lower quality, recently occupied and colder areas arrive later than those from traditionally occupied areas. The population is also expanding into new wintering areas, and males from traditionally occupied winter sites also arrive earlier than those occupying novel sites. 3. Annual variation in timing of migration of individuals is influenced by large-scale weather systems (the North Atlantic Oscillation), but between-individual variation is a stronger predictor of arrival time than the NAO. Distance between winter and breeding sites does not influence arrival times. 4. Annual variation in timing of migration is therefore influenced by climatic factors, but the pattern of individual arrival is primarily related to breeding and winter habitat quality. These habitat effects on arrival patterns are likely to operate through variation in individual condition and local-scale density-dependent processes. Timing of migration thus appears to be a key component of the intricate relationship between wintering and breeding grounds in this migratory system.