The exploitation of biofilm by migrant western sandpipers (Calidrismauri).

Assessing the quality of migratory shorebird stopover sites requires good measures of food availability. We developed simple methods to measure biofilm grazing by migrant western sandpipers (Calidris mauri), a species for which biofilm is an important dietary component. We used a field-portable chlorofluorometer to measure the density of chlorophyll-a (Chl-a) in surficial biofilms on Roberts Bank, a large intertidal mudflat in British Columbia, Canada, during northward migration. Chl-a density begins at a low level during each diurnal emersion period, and increases steadily during emersion at 4.1 mg m-2 h-1 for a total of ∼24.6 mg m-2 over a typical 6 h emersion period and ∼41 mg m-2 over a 10 h emersion period. Western sandpipers grazed at 1.35-1.45 mg Chl-a m-2 min-1, thus biofilm production supports 17.6 min m-2 of grazing time during a 6 h low tide period and 29.3 min m-2 during a 10 h period. During peak northward migration, the average grazing intensity of western sandpipers over an intertidal emersion period was 3.3-6.4 min m-2, suggesting that biofilm accumulation was 2.7-8.8 fold greater than the amount consumed. We found Chl-a density was highest (∼65 mg per m2) within 40 m of the shoreline. Grazing intensity was lowest close to shore, where predation risk from falcon attacks is highest. Grazing intensity peaked at 240 m and then declined, lowering Chl-a density at greater distances to a uniform level of ∼54 mg m-2. These results indicate that interactions between biofilm production and sandpiper grazing underlie spatio-temporal patterns in biofilm abundance on Roberts Bank.

Avian wings can lengthen rather than shorten in response to increased migratory predation danger.

Increasing predation danger can select for safety-enhancing modifications to prey morphology. Here, we document the multi-decade wing lengthening of a Pacific flyway migrant, the western sandpiper (Calidris mauri), and contrast this with contemporaneous wing shortening of the closely related semipalmated sandpiper (C. pusilla) on the Atlantic flyway. We measured >12,000 southbound western sandpipers captured from 1978 to 2020 at a major stopover site in British Columbia. Wing length increased at 0.074 mm year-1 (SE = 0.017; p < .0003) for adults, and 0.087 mm year-1 (SE = 0.029; p < .007) for juveniles. These rates are of similarly large magnitude (4%-5% overall), but opposite in direction, to the rate we previously reported for semipalmated sandpiper adults (-0.103 mm year-1). In both species, the change is specific to wings rather than being part of a general body size change. We interpret both trends as responses to the ongoing strong increase of peregrine falcon (Falco peregrinus) populations since the mid-1970s, an important predator encountered by these species in contrasting ways during migration. Western sandpipers and peregrine migrations have temporal and spatial overlap. Longer wings enhance migratory speed and efficiency, enabling western sandpipers to decrease overlap by advancing to safer zones ahead of falcon passage. In contrast, semipalmated sandpipers primarily encounter peregrines as residents at migratory staging sites. Shorter wings improve acceleration and agility, helping migrants to escape attacks. Juvenile western sandpiper wing length also shows a component additive to the lengthening trend, shifting between years at 0.055 mm day-1 with the highly variable snowmelt date, with wings shorter following early springs. On the Pacific flyway, the timing of peregrine southward passage advances with snowmelt, increasing the relative exposure of juveniles to post-migratory resident peregrines. We interpret this annual wing length adjustment as an induced defense, made possible because snowmelt timing is a reliable cue to danger in the upcoming migration.

Spatial genetic structure of European wild boar, with inferences on late-Pleistocene and Holocene demographic history.

European wildlife has been subjected to intensifying levels of anthropogenic impact throughout the Holocene, yet the main genetic partitioning of many species is thought to still reflect the late-Pleistocene glacial refugia. We analyzed 26,342 nuclear SNPs of 464 wild boar (Sus scrofa) across the European continent to infer demographic history and reassess the genetic consequences of natural and anthropogenic forces. We found that population fragmentation, inbreeding and recent hybridization with domestic pigs have caused the spatial genetic structure to be heterogeneous at the local scale. Underlying local anthropogenic signatures, we found a deep genetic structure in the form of an arch-shaped cline extending from the Dinaric Alps, via Southeastern Europe and the Baltic states, to Western Europe and, finally, to the genetically diverged Iberian peninsula. These findings indicate that, despite considerable anthropogenic influence, the deeper, natural continental structure is still intact. Regarding the glacial refugia, our findings show a weaker signal than generally assumed, but are nevertheless suggestive of two main recolonization routes, with important roles for Southern France and the Balkans. Our results highlight the importance of applying genomic resources and framing genetic results within a species' demographic history and geographic distribution for a better understanding of the complex mixture of underlying processes.

Seasonal and inter-annual variation in exposure to peregrines (Falco peregrinus) for southbound western sandpipers (Calidris mauri).

BACKGROUND: The western sandpiper (Calidris mauri) is an early southbound migrant species in North America. The 'peregrine avoidance' hypothesis proposes that this timing evolved to reduce exposure to their main predator, the peregrine (Falco peregrinus), along the Pacific flyway. METHODS: I evaluate this hypothesis based on 16 years of near-daily (June - October) measures of peregrine presence made on the Fraser River estuary, a major stopover in the Pacific northwest. RESULTS: Exposure to peregrines is lowest for the earliest southbound western sandpipers, and rises steeply as peregrines en route from northern breeding areas begin to arrive in late July or August. Peregrine arrival timing varies greatly between years, shifting in step with the onset of spring along coastal Alaska. Peregrine presence on the Fraser estuary on any date is higher in years with earlier spring onset. On the median adult sandpiper passage date (day-of-year 198) this increases 17-fold over the inter-annual range between the earliest and latest peregrine arrival dates. CONCLUSION: The pattern of strong and predictable changes in the seasonal pattern of danger quantified here provides a further test of the hypothesis that danger affects migratory timing. Western sandpipers appear to anticipate the exposure level of southward migration, perhaps because they are able to observe spring onset on their Alaskan breeding grounds. They adjust the duration of parental care and length of the breeding season to keep the date of migratory departure from the Arctic relatively invariant in spite of large interannual variation in spring onset. While underway they also adjust aspects of migratory behavior. These observations support the 'peregrine avoidance' hypothesis, and suggest that western sandpipers are able to counter, at least partially, the higher migratory danger of early spring years.

Predation landscapes influence migratory prey ecology and evolution.

Migratory prey experience spatially variable predation across their life cycle. They face unique challenges in navigating this predation landscape, which affects their perception of risk, antipredator responses, and resulting mortality. Variable and unfamiliar predator cues during migration can limit accurate perception of risk and migrants often rely on social information and learning to compensate. The energetic demands of migration constrain antipredator responses, often through context-dependent patterns. While migration can increase mortality, migrants employ diverse strategies to balance risks and rewards, including life history and antipredator responses. Humans interact frequently with migratory prey across space and alter both mortality risk and antipredator responses, which can scale up to affect migratory populations and should be considered in conservation and management.

Oversummering juvenile and adult Semipalmated sandpipers in Perú gain enough survival to compensate for foregone breeding opportunity.

BACKGROUND: Age at maturity and the timing of first breeding are important life history traits. Most small shorebird species mature and breed as 'yearlings', but have lower reproductive success than adults. In some species, yearlings may defer northward migration and remain in non-breeding regions ('oversummering') until they reach 2 years of age. Some adults also oversummer. Oversummering would be favoured by natural selection if survival were as a result raised sufficiently to compensate for the missed breeding opportunity. Several thousand Semipalmated Sandpipers (Calidris pusilla) spend the non-breeding period at Paracas, Perú, including individuals with long bills (likely from eastern Arctic breeding populations ~ 8000 km distant) and short bills (likely from western Arctic breeding populations, up to 11,000 km distant), with short-billed birds more likely to oversummer. We tested the prediction that oversummering birds have higher survival than migrants, and that the magnitude of this higher survival for oversummering birds is enough to compensate for their lost breeding season. METHODS: We used a Multi-State Mark-Recapture model based on 5 years of encounter data (n = 1963 marked birds, and 3229 resightings) obtained year-round at Paracas, Perú, to estimate seasonal (i.e. breeding and non-breeding) survivorship for migrant and oversummering birds. We calculated the magnitude of the oversummering survival advantage required to compensate, for both yearlings and adults, based on published measures of annual survival and reproductive success. Using bill length as a proxy for migration distance, we investigated whether migratory survival is distance-dependent. RESULTS: We estimate that 28% of yearlings and 19% of adults oversummer. Survival is higher for oversummering birds than for migrants, and the oversummering survival advantage is greater for adults (0.215) than for yearlings (0.140). The theoretical thresholds predicted by the size of the missed reproductive opportunity are 0.240 for adults and 0.134 for yearlings. Migratory survival decreases and the oversummering rate increases with migration distance, as assessed by culmen length. CONCLUSIONS: Our results support the life history hypothesis that oversummering raises survival enough to compensate for the loss of a breeding opportunity. Greater migration distance lowers survival and increases the probability of oversummering.

A history of hybrids? Genomic patterns of introgression in the True Geese.

BACKGROUND: The impacts of hybridization on the process of speciation are manifold, leading to distinct patterns across the genome. Genetic differentiation accumulates in certain genomic regions, while divergence is hampered in other regions by homogenizing gene flow, resulting in a heterogeneous genomic landscape. A consequence of this heterogeneity is that genomes are mosaics of different gene histories that can be compared to unravel complex speciation and hybridization events. However, incomplete lineage sorting (often the outcome of rapid speciation) can result in similar patterns. New statistical techniques, such as the D-statistic and hybridization networks, can be applied to disentangle the contributions of hybridization and incomplete lineage sorting. We unravel patterns of hybridization and incomplete lineage sorting during and after the diversification of the True Geese (family Anatidae, tribe Anserini, genera Anser and Branta) using an exon-based hybridization network approach and taking advantage of discordant gene tree histories by re-sequencing all taxa of this clade. In addition, we determine the timing of introgression and reconstruct historical effective population sizes for all goose species to infer which demographic or biogeographic factors might explain the observed patterns of introgression. RESULTS: We find indications for ancient interspecific gene flow during the diversification of the True Geese and were able to pinpoint several putative hybridization events. Specifically, in the genus Branta, both the ancestor of the White-cheeked Geese (Hawaiian Goose, Canada Goose, Cackling Goose and Barnacle Goose) and the ancestor of the Brent Goose hybridized with Red-breasted Goose. One hybridization network suggests a hybrid origin for the Red-breasted Goose, but this scenario seems unlikely and it not supported by the D-statistic analysis. The complex, highly reticulated evolutionary history of the genus Anser hampered the estimation of ancient hybridization events by means of hybridization networks. The reconstruction of historical effective population sizes shows that most species showed a steady increase during the Pliocene and Pleistocene. These large effective population sizes might have facilitated contact between diverging goose species, resulting in the establishment of hybrid zones and consequent gene flow. CONCLUSIONS: Our analyses suggest that the evolutionary history of the True Geese is influenced by introgressive hybridization. The approach that we have used, based on genome-wide phylogenetic incongruence and network analyses, will be a useful procedure to reconstruct the complex evolutionary histories of many naturally hybridizing species groups.

Long-term continental changes in wing length, but not bill length, of a long-distance migratory shorebird.

We compiled a >50-year record of morphometrics for semipalmated sandpipers (Calidris pusilla), a shorebird species with a Nearctic breeding distribution and intercontinental migration to South America. Our data included >57,000 individuals captured 1972-2015 at five breeding locations and three major stopover sites, plus 139 museum specimens collected in earlier decades. Wing length increased by ca. 1.5 mm (>1%) prior to 1980, followed by a decrease of 3.85 mm (nearly 4%) over the subsequent 35 years. This can account for previously reported changes in metrics at a migratory stopover site from 1985 to 2006. Wing length decreased at a rate of 1,098 darwins, or 0.176 haldanes, within the ranges of other field studies of phenotypic change. Bill length, in contrast, showed no consistent change over the full period of our study. Decreased body size as a universal response of animal populations to climate warming, and several other potential mechanisms, are unable to account for the increasing and decreasing wing length pattern observed. We propose that the post-WWII near-extirpation of falcon populations and their post-1973 recovery driven by the widespread use and subsequent limitation on DDT in North America selected initially for greater flight efficiency and latterly for greater agility. This predation danger hypothesis accounts for many features of the morphometric data and deserves further investigation in this and other species.

The redistribution of non-breeding dunlins in response to the post-DDT recovery of falcons.

Large-scale changes in predator populations are occurring worldwide due to (re-)introductions, over-exploitation, or recovery after decimation by pesticides and persecution. These widespread changes may affect the distribution of their prey. We studied the continental-scale distributions of non-breeding Calidris alpina pacifica and C. a. hudsonia (Pacific and Atlantic dunlins, respectively), as numbers of their major predators-peregrines (Falco peregrinus) and merlins (F. columbarius; together 'falcons')-increased after DDT was banned in 1973. For the period 1975-2010 we compiled the number of dunlins and falcons in each of 244 Christmas Bird Count circles, which cover most of the dunlins' non-breeding ranges. Over the study period, falcons increased by 6.5- (Pacific) and 3.1- (Atlantic) fold, spread to more count circles, and the number of dunlins per falcon fell. The annual total count of the two dunlin sub-species fluctuated strongly and independently. We measured annual aggregation as the expected proportion of a subspecies total found on the same count circle as a randomly selected dunlin. The average aggregation of Pacific dunlins (0.117) was about double that of Atlantic dunlins (0.059), with annual variation largely attributable to changes in a few large count circles. The slope of the aggregative response to year-on-year changes in dunlin numbers was identical on the two coasts. The response to the ongoing falcon increase was positive and significant on the Pacific and slightly negative on the Atlantic. We interpret these results using a version of the ideal free distribution that includes predation danger.

Spatial variation in polycyclic aromatic hydrocarbon exposure in Barrow's goldeneye (Bucephala islandica) in coastal British Columbia.

Barrow's goldeneyes are sea ducks that winter throughout coastal British Columbia (BC). Their diet consists primarily of intertidal blue mussels, which can accumulate PAHs; accordingly, goldeneyes may be susceptible to exposure through contaminated prey. In 2014/15, we examined total PAH concentrations in mussels from undeveloped and developed coastal areas of BC. At those same sites, we used EROD to measure hepatic CYP1A induction in goldeneyes. We found higher mussel PAH concentrations at developed coastal sites. Regionally, goldeneyes from southern BC, which has relatively higher coastal development, had higher EROD activity compared to birds from northern BC. Our results suggest goldeneyes wintering in coastal BC were exposed to PAHs through diet, with higher exposure among birds wintering in coastal areas with greater anthropogenic influence. These results suggest the mussel-goldeneye system is suitable as a natural, multi-trophic-level indicator of contemporary hydrocarbon contamination occurrence and exposure useful for establishing oil spill recovery endpoints.

A tree of geese: A phylogenomic perspective on the evolutionary history of True Geese.

Phylogenetic incongruence can be caused by analytical shortcomings or can be the result of biological processes, such as hybridization, incomplete lineage sorting and gene duplication. Differentiation between these causes of incongruence is essential to unravel complex speciation and diversification events. The phylogeny of the True Geese (tribe Anserini, Anatidae, Anseriformes) was, until now, contentious, i.e., the phylogenetic relationships and the timing of divergence between the different goose species could not be fully resolved. We sequenced nineteen goose genomes (representing seventeen species of which three subspecies of the Brent Goose, Branta bernicla) and used an exon-based phylogenomic approach (41,736 exons, representing 5887 genes) to unravel the evolutionary history of this bird group. We thereby provide general guidance on the combination of whole genome evolutionary analyses and analytical tools for such cases where previous attempts to resolve the phylogenetic history of several taxa could not be unravelled. Identical topologies were obtained using either a concatenation (based upon an alignment of 6,630,626 base pairs) or a coalescent-based consensus method. Two major lineages, corresponding to the genera Anser and Branta, were strongly supported. Within the Branta lineage, the White-cheeked Geese form a well-supported sub-lineage that is sister to the Red-breasted Goose (Branta ruficollis). In addition, two main clades of Anser species could be identified, the White Geese and the Grey Geese. The results from the consensus method suggest that the diversification of the genus Anser is heavily influenced by rapid speciation and by hybridization, which may explain the failure of previous studies to resolve the phylogenetic relationships within this genus. The majority of speciation events took place in the late Pliocene and early Pleistocene (between 4 and 2millionyears ago), conceivably driven by a global cooling trend that led to the establishment of a circumpolar tundra belt and the emergence of temperate grasslands. Our approach will be a fruitful strategy for resolving many other complex evolutionary histories at the level of genera, species, and subspecies.

Hybridization in geese: a review.

The high incidence of hybridization in waterfowl (ducks, geese and swans) makes this bird group an excellent study system to answer questions related to the evolution and maintenance of species boundaries. However, knowledge on waterfowl hybridization is biased towards ducks, with a large knowledge gap in geese. In this review, we assemble the available information on hybrid geese by focusing on three main themes: (1) incidence and frequency, (2) behavioural mechanisms leading to hybridization, and (3) hybrid fertility. Hybridization in geese is common on a species-level, but rare on a per-individual level. An overview of the different behavioural mechanisms indicates that forced extra-pair copulations and interspecific nest parasisitm can both lead to hybridization. Other sources of hybrids include hybridization in captivity and vagrant geese, which may both lead to a scarcity of conspecifics. The different mechanisms are not mutually exclusive and it is currently not possible to discriminate between the different mechanisms without quantitative data. Most hybrid geese are fertile; only in crosses between distantly related species do female hybrids become sterile. This fertility pattern, which is in line with Haldane's Rule, may facilitate interspecific gene flow between closely related species. The knowledge on hybrid geese should be used, in combination with the information available on hybridization in ducks, to study the process of avian speciation.

Goal-dependent current compensation and drift in surf scoter flocks.

BACKGROUND: Animals moving through air or water toward a goal frequently must contend with fluid currents, which can drift the actual path of the animal away from the direction of heading. Whether, and to what degree, animals compensate for currents depends on the species and environmental context, but plays an important role in the movement ecology of the species. In this paper, flocks of surf scoters (Melanitta perspicillata), an aquatic diving duck, were individually tracked during collective foraging in the presence of sideward water currents to assess the individual compensatory response while moving from open water toward the foraging location versus return to open water. RESULTS: During short-range movement toward the foraging location, surf scoters moved more slowly, and compensated for currents by orienting diagonally into the current to maintain a perpendicular track to the goal. In contrast, during return to open water, surf scoters moved faster, and maintained a perpendicular orientation away from the foraging location, and allowed the sideward current to drift their track diagonally. CONCLUSIONS: Surf scoters show a behavioural flexibility in response to currents, alternately using compensation and drift as the movement goal and consequent cost of accuracy change.

Do Arctic breeding geese track or overtake a green wave during spring migration?

Geese breeding in the Arctic have to do so in a short time-window while having sufficient body reserves. Hence, arrival time and body condition upon arrival largely influence breeding success. The green wave hypothesis posits that geese track a successively delayed spring flush of plant development on the way to their breeding sites. The green wave has been interpreted as representing either the onset of spring or the peak in nutrient biomass. However, geese tend to adopt a partial capital breeding strategy and might overtake the green wave to accomplish a timely arrival on the breeding site. To test the green wave hypothesis, we link the satellite-derived onset of spring and peak in nutrient biomass with the stopover schedule of individual Barnacle Geese. We find that geese track neither the onset of spring nor the peak in nutrient biomass. Rather, they arrive at the southernmost stopover site around the peak in nutrient biomass, and gradually overtake the green wave to match their arrival at the breeding site with the local onset of spring, thereby ensuring gosling benefit from the peak in nutrient biomass. Our approach for estimating plant development stages is critical in testing the migration strategies of migratory herbivores.

Global lack of flyway structure in a cosmopolitan bird revealed by a genome wide survey of single nucleotide polymorphisms.

Knowledge about population structure and connectivity of waterfowl species, especially mallards (Anas platyrhynchos), is a priority because of recent outbreaks of avian influenza. Ringing studies that trace large-scale movement patterns have to date been unable to detect clearly delineated mallard populations. We employed 363 single nucleotide polymorphism markers in combination with population genetics and phylogeographical approaches to conduct a population genomic test of panmixia in 801 mallards from 45 locations worldwide. Basic population genetic and phylogenetic methods suggest no or very little population structure on continental scales. Nor could individual-based structuring algorithms discern geographical structuring. Model-based coalescent analyses for testing models of population structure pointed to strong genetic connectivity among the world's mallard population. These diverse approaches all support the conclusion that there is a lack of clear population structure, suggesting that the world's mallards, perhaps with minor exceptions, form a single large, mainly interbreeding population.

Widespread horizontal genomic exchange does not erode species barriers among sympatric ducks.

BACKGROUND: The study of speciation and maintenance of species barriers is at the core of evolutionary biology. During speciation the genome of one population becomes separated from other populations of the same species, which may lead to genomic incompatibility with time. This separation is complete when no fertile offspring is produced from inter-population matings, which is the basis of the biological species concept. Birds, in particular ducks, are recognised as a challenging and illustrative group of higher vertebrates for speciation studies. There are many sympatric and ecologically similar duck species, among which fertile hybrids occur relatively frequently in nature, yet these species remain distinct. RESULTS: We show that the degree of shared single nucleotide polymorphisms (SNPs) between five species of dabbling ducks (genus Anas) is an order of magnitude higher than that previously reported between any pair of eukaryotic species with comparable evolutionary distances. We demonstrate that hybridisation has led to sustained exchange of genetic material between duck species on an evolutionary time scale without disintegrating species boundaries. Even though behavioural, genetic and ecological factors uphold species boundaries in ducks, we detect opposing forces allowing for viable interspecific hybrids, with long-term evolutionary implications. Based on the superspecies concept we here introduce the novel term "supra-population" to explain the persistence of SNPs identical by descent within the studied ducks despite their history as distinct species dating back millions of years. CONCLUSIONS: By reviewing evidence from speciation theory, palaeogeography and palaeontology we propose a fundamentally new model of speciation to accommodate our genetic findings in dabbling ducks. This model, we argue, may also shed light on longstanding unresolved general speciation and hybridisation patterns in higher organisms, e.g. in other bird groups with unusually high hybridisation rates. Observed parallels to horizontal gene transfer in bacteria facilitate the understanding of why ducks have been such an evolutionarily successful group of animals. There is large evolutionary potential in the ability to exchange genes among species and the resulting dramatic increase of effective population size to counter selective constraints.

Variable and complex food web structures revealed by exploring missing trophic links between birds and biofilm.

Food webs are comprised of a network of trophic interactions and are essential to elucidating ecosystem processes and functions. However, the presence of unknown, but critical networks hampers understanding of complex and dynamic food webs in nature. Here, we empirically demonstrate a missing link, both critical and variable, by revealing that direct predator-prey relationships between shorebirds and biofilm are widespread and mediated by multiple ecological and evolutionary determinants. Food source mixing models and energy budget estimates indicate that the strength of the missing linkage is dependent on predator traits (body mass and foraging action rate) and the environment that determines food density. Morphological analyses, showing that smaller bodied species possess more developed feeding apparatus to consume biofilm, suggest that the linkage is also phylogenetically dependent and affords a compelling re-interpretation of niche differentiation. We contend that exploring missing links is a necessity for revealing true network structure and dynamics.

Evolution and connectivity in the world-wide migration system of the mallard: inferences from mitochondrial DNA.

BACKGROUND: Main waterfowl migration systems are well understood through ringing activities. However, in mallards (Anas platyrhynchos) ringing studies suggest deviations from general migratory trends and traditions in waterfowl. Furthermore, surprisingly little is known about the population genetic structure of mallards, and studying it may yield insight into the spread of diseases such as Avian Influenza, and in management and conservation of wetlands. The study of evolution of genetic diversity and subsequent partitioning thereof during the last glaciation adds to ongoing discussions on the general evolution of waterfowl populations and flyway evolution. Hypothesised mallard flyways are tested explicitly by analysing mitochondrial mallard DNA from the whole northern hemisphere. RESULTS: Phylogenetic analyses confirm two mitochondrial mallard clades. Genetic differentiation within Eurasia and North-America is low, on a continental scale, but large differences occur between these two land masses (F(ST) = 0.51). Half the genetic variance lies within sampling locations, and a negligible portion between currently recognised waterfowl flyways, within Eurasia and North-America. Analysis of molecular variance (AMOVA) at continent scale, incorporating sampling localities as smallest units, also shows the absence of population structure on the flyway level. Finally, demographic modelling by coalescence simulation proposes a split between Eurasia and North-America 43,000 to 74,000 years ago and strong population growth (~100fold) since then and little migration (not statistically different from zero). CONCLUSIONS: Based on this first complete assessment of the mallard's world-wide population genetic structure we confirm that no more than two mtDNA clades exist. Clade A is characteristic for Eurasia, and clade B for North-America although some representatives of clade A are also found in North-America. We explain this pattern by evaluating competing hypotheses and conclude that a complex mix of historical, recent and anthropogenic factors shaped the current mallard populations. We refute population classification based on flyways proposed by ornithologists and managers, because they seem to have little biological meaning. Our results have implications for wetland management and conservation, with special regard to the release of farmed mallards for hunting, as well as for the possible transmission of Avian Influenza by mallards due to migration.

Avian influenza surveillance with FTA cards: field methods, biosafety, and transportation issues solved.

Avian Influenza Viruses (AIVs) infect many mammals, including humans(1). These AIVs are diverse in their natural hosts, harboring almost all possible viral subtypes(2). Human pandemics of flu originally stem from AIVs(3). Many fatal human cases during the H5N1 outbreaks in recent years were reported. Lately, a new AIV related strain swept through the human population, causing the 'swine flu epidemic'(4). Although human trading and transportation activity seems to be responsible for the spread of highly pathogenic strains(5), dispersal can also partly be attributed to wild birds(6, 7). However, the actual reservoir of all AIV strains is wild birds. In reaction to this and in face of severe commercial losses in the poultry industry, large surveillance programs have been implemented globally to collect information on the ecology of AIVs, and to install early warning systems to detect certain highly pathogenic strains(8-12). Traditional virological methods require viruses to be intact and cultivated before analysis. This necessitates strict cold chains with deep freezers and heavy biosafety procedures to be in place during transport. Long-term surveillance is therefore usually restricted to a few field stations close to well equipped laboratories. Remote areas cannot be sampled unless logistically cumbersome procedures are implemented. These problems have been recognised(13, 14) and the use of alternative storage and transport strategies investigated (alcohols or guanidine)(15-17). Recently, Kraus et al.(18) introduced a method to collect, store and transport AIV samples, based on a special filter paper. FTA cards(19) preserve RNA on a dry storage basis(20) and render pathogens inactive upon contact(21). This study showed that FTA cards can be used to detect AIV RNA in reverse-transcription PCR and that the resulting cDNA could be sequenced and virus genes and determined. In the study of Kraus et al.(18) a laboratory isolate of AIV was used, and samples were handled individually. In the extension presented here, faecal samples from wild birds from the duck trap at the Ottenby Bird Observatory (SE Sweden) were tested directly to illustrate the usefulness of the methods under field conditions. Catching of ducks and sample collection by cloacal swabs is demonstrated. The current protocol includes up-scaling of the work flow from single tube handling to a 96-well design. Although less sensitive than the traditional methods, the method of FTA cards provides an excellent supplement to large surveillance schemes. It allows collection and analysis of samples from anywhere in the world, without the need to maintaining a cool chain or safety regulations with respect to shipping of hazardous reagents, such as alcohol or guanidine.

Interactions between rate processes with different timescales explain counterintuitive foraging patterns of arctic wintering eiders.

To maximize fitness, animals must respond to a variety of processes that operate at different rates or timescales. Appropriate decisions could therefore involve complex interactions among these processes. For example, eiders wintering in the arctic sea ice must consider locomotion and physiology of diving for benthic invertebrates, digestive processing rate and a nonlinear decrease in profitability of diving as currents increase over the tidal cycle. Using a multi-scale dynamic modelling approach and continuous field observations of individuals, we demonstrate that the strategy that maximizes long-term energy gain involves resting during the most profitable foraging period (slack currents). These counterintuitive foraging patterns are an adaptive trade-off between multiple overlapping rate processes and cannot be explained by classical rate-maximizing optimization theory, which only considers a single timescale and predicts a constant rate of foraging. By reducing foraging and instead digesting during slack currents, eiders structure their activity in order to maximize long-term energetic gain over an entire tide cycle. This study reveals how counterintuitive patterns and a complex functional response can result from a simple trade-off among several overlapping rate processes, emphasizing the necessity of a multi-scale approach for understanding adaptive routines in the wild and evaluating mechanisms in ecological time series.