The role of phenolic compounds and nutrients in determining food preference in greater snow geese

Summary We tested Buchsbaum's hypothesis that food palatability in geese is determined by a hierarchy of feeding cues among which deterrent secondary metabolites (mostly phenols) have a primary role (Buchsbaum et al. 1984). In preference tests, greater snow goose feeding was slightly depressed when grass was sprayed with ferulic acid but not when grass was sprayed with p-coumaric and tannic acids. Extracts of Timothy grass, red clover or alfalfa sprayed on grass also failed to depress goose feeding. In a multifactor experiment, phenol and protein content and height of grass were manipulated simultaneously. When ferulic acid was sprayed, protein and phenol content interacted in determining goose feeding preferences; protein content had no effect in the absence of phenol but did have an effect when phenol was added. When tannic acid was used in a similar experiment, results were inconclusive because of a significant and complex interaction between protein content and height of grass. Our results generally failed to support Buchsbaum's hypothesis that phenol content of plants has a primary role in determining food preference in geese. Protein content of plants seemed to be a more important factor.     

Energetics of reproduction in female and male greater snow geese

Arctic-nesting geese are classified as capital breeders (i.e., birds that rely largely on endogenous reserves to meet the high nutrient requirement of clutch formation) as opposed to income breeders (those that rely directly on ingested food). However, some evidence has suggested that energy reserves of greater snow geese (Chen caerulescens atlantica) are not sufficient to meet these costs. We tested whether greater snow geese were capital breeders and examined the consequences on their reproductive strategy. We collected 48 females and 47 males from arrival at the breeding colony on Bylot Island (NWT) to the postlaying stage, and determined fat and protein content in somatic and reproductive tissues. Rapid follicular growth was initiated only after arrival on the breeding ground. Somatic fat of females tended to increase during the prelaying and laying stages at the same time that most of the fat was deposited in developing follicles. Decrease in somatic fat in postlaying females was mostly related to its use for meeting metabolic requirements during early incubation. Hence, almost all fat invested in the clutch came from food intake, not endogenous reserves. Somatic protein was maintained during prelaying but decreased during laying, suggesting that some protein reserves were deposited in the eggs. There was no relationship between somatic fat and the number of developing follicles (incubating females excluded) but a relationship was found with somatic protein. Clutch size was not related to body size. In males, somatic fat, which was similar to females at arrival, was almost completely depleted by the postlaying stage. Male somatic protein remained stable. Male somatic reserves were not related to the somatic reserves, clutch size or nutrient investment in reproduction of their mate. We conclude that, in female greater snow geese, little fat reserves (if any) were used for egg formation though some protein reserves were used, whereas males relied heavily on their fat reserves during the same period. This is probably a consequence of the long and costly migration between the last staging area and the breeding grounds.     

Effects of fleas on nest success of Arctic barnacle geese: experimentally testing the mechanism

Parasites have detrimental effects on their hosts’ fitness. Therefore, behavioural adaptations have evolved to avoid parasites or, when an individual is already in contact with a parasite, prevent or minimize infections. Such anti‐parasite behaviours can be very effective, but can also be costly for the host. Specifically, ectoparasites can elicit strong host anti‐parasite behaviours and interactions between fleas (Siphonaptera) and their hosts are one of the best studied. In altricial bird species, nest fleas can negatively affect both parent and offspring fitness components. However, knowledge on the effects of fleas on precocial bird species is scarce. Research on geese in the Canadian Arctic indicated that fleas have a negative impact on reproductive success. One possible hypothesis is that fleas may affect female incubation behaviour. Breeding females with many fleas in their nest may increase the frequency and/or duration of incubation breaks and could even totally desert their nest. The aim of our study was to 1) determine if a similar negative relationship existed between flea abundance and reproductive success in our study colony of Arctic breeding barnacle geese Branta leucopsis and 2) experimentally quantify if such effects could be explained by a negative effect of nest fleas on female behaviour. We compared host anti‐parasite and incubation behaviour between experimentally flea‐reduced and control nests using wildlife cameras and temperature loggers. We found that flea abundance was negatively associated with hatching success. We found little experimental support, however, for changes in behaviour of the breeding female as a possible mechanism to explain this effect.     

The dynamics of landscape change and snow geese in mid-continent North America

The Mid‐Continent Population of the lesser snow goose, which breeds in the eastern and central Canadian Arctic and sub‐Arctic, and winters in the southern United States and northern Mexico has increased 5–7% annually from the late 1960s to the mid‐1990s, largely because of increased survival in response to an agricultural food subsidy. The rise in numbers complements the increased use of nitrogen fertilizers and a corresponding rise in yields of rice, corn, and wheat along the flyways and on the wintering grounds. In sub‐Arctic migration areas and at Arctic breeding colonies, foraging by high numbers of birds has led to loss of coastal vegetation, adverse changes in soil properties and the establishment of an alternative stable state of exposed sediment, which can be detected with LANDSAT imagery. At a local scale, gosling growth, size and survival decreased in affected areas and other taxa have been adversely affected. The food subsidy on wintering and migration areas appears insufficient to meet reproductive demands as foraging in spring continues to occur on southern Hudson Bay staging and nesting areas. The recent introduction of liberal hunting regulations may reduce population size in the near term, but the revegetation of these coastal ecosystems will take decades to achieve. The present pattern of vegetation loss in these Arctic coastal systems is likely to continue in the forseeable future.     

Comparative survival and recovery of Ross's and lesser snow geese from Canada's central arctic

Large increases in several populations of North American arctic geese have resulted in ecosystem-level effects from associated herbivory. Consequently, some breeding populations have shown density dependence in recruitment through declines in food availability. Differences in population trajectories of lesser snow geese (Chen caerulescens caerulescens; hereafter snow geese) and Ross's geese (C. rossii) breeding in mixed-species colonies south of Queen Maud Gulf (QMG), in Canada's central arctic, suggest that density dependence may be limiting snow goose populations. Specifically, long-term declines in age ratios (immature:adult) of harvested snow geese may have resulted from declines in juvenile survival. Thus, we focused on juvenile (first-year) survival of snow and Ross's geese in relation to timing of reproduction (annual mean nest initiation date) and late summer weather. We banded Ross's and snow geese from 1991 to 2008 in the QMG Migratory Bird Sanctuary. We used age-structured mark-recapture models to estimate annual survival rates for adults and juveniles from recoveries of dead birds. Consistent with life history differences, juvenile snow geese survived at rates higher than juvenile Ross's geese. Juvenile survival of both species also was lower in late seasons, but was unrelated to arctic weather measured during a 17-day period after banding. We found no evidence of density dependence (i.e., a decline in juvenile survival over time) in either species. We also found no interspecific differences in age-specific hunting vulnerability, though juveniles were more vulnerable than adults in both species, as expected. Thus, interspecific differences in survival were unrelated to harvest. Lower survival of juvenile Ross's geese may result from natural migration mortality related to smaller body size (e.g., greater susceptibility to inclement weather or predation) compared to juvenile snow geese. Despite lower first-year survival, recruitment by Ross's geese may still be greater than that by snow geese because of earlier sexual maturity, greater breeding propensity, and higher nest success by Ross's geese. © 2012 The Wildlife Society.     

Rapid formation of new migration route and breeding area by Arctic geese

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Intra- and interspecific differences in palatability of Arctic macroalgae from Kongsfjorden (Spitsbergen) for two benthic sympatric invertebrates

Information on herbivore-macroalgae interactions is extremely scarce for Arctic habitats. In this study, the potential of 19 macroalgae as food source for herbivores was investigated for the first time in Arctic waters (Kongsfjorden, Spitsbergen) with emphasis on algal defense against grazing. Only two of the 19 tested macroalgae-associated invertebrates consumed macroalgae in measurable amounts, the locally abundant green sea urchin Strongylocentrotus droebachiensis (OF Müller) and the amphipod Gammarellus homari (JC Fabricius). In the inner basin of the fjord, the dense macroalgal stock represents a potential food source. However, in this area, herbivory plays only a minor role. In contrast, in the outer basin of Kongsfjorden S. droebachiensis exerts a strong top-down control on macroalgal assemblages.Laboratory feeding assays in the with the two herbivores showed grazer-specific feeding preferences. The amphipod G. homari exhibits a preference for delicate red algal species like Devaleraea ramentacea, whereas the sea urchin S. droebachiensis significantly preferred more leathery seaweeds like Laminaria and Alaria. The red alga Palmaria palmata is a very attractive food for both herbivores, while the brown alga Desmarestia viridis and the red alga Ptilota gunneri are among the least preferred algae. To distinguish between physical and tissue-specific plant properties, which have a deterring or stimulating effect on the grazing behaviour, both grazers were offered intact algal tissue and artificial food in separate feeding assays. While physical and tissue-specific plant properties-especially in the Laminariales-deterred G. homari, these properties did not deter S. droebachiensis.This study provides insights into herbivore consumption in the benthic food web of Kongsfjorden. As a general potential top-down factor controlling primary production, herbivory needs to be quantified further in Kongsfjorden to develop adequate carbon flow models for this important reference site for the study of climate change on high latitude marine ecosystems.     

Winter precipitation and snow accumulation drive the methane sink or source strength of Arctic tussock tundra

Arctic winter precipitation is projected to increase with global warming, but some areas will experience decreases in snow accumulation. Although Arctic CH₄ emissions may represent a significant climate forcing feedback, long‐term impacts of changes in snow accumulation on CH₄ fluxes remain uncertain. We measured ecosystem CH₄ fluxes and soil CH₄ and CO₂ concentrations and ¹³C composition to investigate the metabolic pathways and transport mechanisms driving moist acidic tundra CH₄ flux over the growing season (Jun–Aug) after 18 years of experimental snow depth increases and decreases. Deeper snow increased soil wetness and warming, reducing soil %O₂ levels and increasing thaw depth. Soil moisture, through changes in soil %O₂ saturation, determined predominance of methanotrophy or methanogenesis, with soil temperature regulating the ecosystem CH₄ sink or source strength. Reduced snow (RS) increased the fraction of oxidized CH₄ (Fox) by 75–120% compared to Ambient, switching the system from a small source to a net CH₄ sink (21 ± 2 and ?31 ± 1 mg CH₄ m^?2 season^?1 at Ambient and RS). Deeper snow reduced Fox by 35–40% and 90–100% in medium‐ (MS) and high‐ (HS) snow additions relative to Ambient, contributing to increasing the CH₄ source strength of moist acidic tundra (464 ± 15 and 3561 ± 97 mg CH₄ m^?2 season^?1 at MS and HS). Decreases in Fox with deeper snow were partly due to increases in plant‐mediated CH₄ transport associated with the expansion of tall graminoids. Deeper snow enhanced CH₄ production within newly thawed soils, responding mainly to soil warming rather than to increases in acetate fermentation expected from thaw‐induced increases in SOC availability. Our results suggest that increased winter precipitation will increase the CH₄ source strength of Arctic tundra, but the resulting positive feedback on climate change will depend on the balance between areas with more or less snow accumulation than they are currently facing.     

Seasonal variation in growth of greater snow goose goslings: the role of food supply

Even though growth rate is an important fitness component, it is still controversial to what extent parent birds adjust the timing of offspring hatch to natural variations in food supply to maximize offspring growth. We studied the role of food availability in explaining inter- and intra-seasonal variation of growth rate in goslings of greater snow geese over 5 years. The peak of hatching generally coincided with the peak of food availability. However, early-hatched goslings usually grew faster than birds hatched at the peak, which in␣turn grew faster than late-hatched goslings, although this phenomenon was not observed in all years. There was considerable variation in growth rate among the five years, the smallest goslings produced in the best year (1991) being larger than the largest goslings of the poorest year (1994). We developed three indices of food availability, based on the cumulative availability of plant biomass and nitrogen content during the growth period, and showed that the cumulative exposure to nitrogen biomass explained up to 43% of variation (intra- and inter-annual) in body size just before fledging. In years with good feeding conditions, early-hatched goslings had access to more nitrogen during their growing period than those hatching on or after the peak and they grew faster. In years of lower food availability, early-hatched goslings had no detectable advantage over peak- or late-hatched birds for access to protein-rich food and no seasonal decline in growth rate was observed. These results confirm the critical role of food supply in the seasonal variation of growth rate in Arctic-nesting geese. Received: 27 June 1997 / Accepted: 9 October 1997     

Isoenzyme status and genetic variability of serum esterases in the lesser snow goose,Anser caerulescens caerulescens

A maximum of 22 bands comprising four esterase subgroups—acetylesterase, carboxylesterase, cholinesterase, and acetylcholinesterase—were detected following electrophoresis of lesser snow goose sera on polyacrylamide gels. A minimum of seven structural genes was surmised to be involved in the biosynthesis of these enzymes following physiochemical characterizations. The genetic variability of these loci was calculated to be 1.25% average heterozygosity, while 14.3% of the loci were polymorphic. These estimates of genetic variability were substantially lower than those reported for other vertebrate species. The low degree of genetic variability found in snow goose serum esterases coupled with the extensive protein multiplicity observed may possibly reflect an adaptive strategy based on biochemical plasticity rather than genic heterozygosity for this species. The nature of evolutionary forces acting upon multiple enzyme systems such as esterases is discussed. The concept of conditional neutrality is introduced and defined within this context.This research was carried out under grants from the National Research Council of Canada and the Canadian Wildlife Service to F. Cooke. J. Grossfield was supported by grants from the National Institutes of Health, GM 21630 and FRAP 10576.     

Contrasting consequences of climate change for migratory geese: Predation,density dependence and carryover effects offset benefits of high‐arctic warming

Climate change is most rapid in the Arctic, posing both benefits and challenges for migratory herbivores. However, population‐dynamic responses to climate change are generally difficult to predict, due to concurrent changes in other trophic levels. Migratory species are also exposed to contrasting climate trends and density regimes over the annual cycle. Thus, determining how climate change impacts their population dynamics requires an understanding of how weather directly or indirectly (through trophic interactions and carryover effects) affects reproduction and survival across migratory stages, while accounting for density dependence. Here, we analyse the overall implications of climate change for a local non‐hunted population of high‐arctic Svalbard barnacle geese, Branta leucopsis, using 28 years of individual‐based data. By identifying the main drivers of reproductive stages (egg production, hatching and fledging) and age‐specific survival rates, we quantify their impact on population growth. Recent climate change in Svalbard enhanced egg production and hatching success through positive effects of advanced spring onset (snow melt) and warmer summers (i.e. earlier vegetation green‐up) respectively. Contrastingly, there was a strong temporal decline in fledging probability due to increased local abundance of the Arctic fox, the main predator. While weather during the non‐breeding season influenced geese through a positive effect of temperature (UK wintering grounds) on adult survival and a positive carryover effect of rainfall (spring stopover site in Norway) on egg production, these covariates showed no temporal trends. However, density‐dependent effects occurred throughout the annual cycle, and the steadily increasing total flyway population size caused negative trends in overwinter survival and carryover effects on egg production. The combination of density‐dependent processes and direct and indirect climate change effects across life history stages appeared to stabilize local population size. Our study emphasizes the need for holistic approaches when studying population‐dynamic responses to global change in migratory species.     

Forage digestibility and intake by lesser snow geese: effects of dominance and resource heterogeneity

We measured forage intake, digestibility, and retention time for 11 free-ranging, human-imprinted lesser snow geese (Chen caerulescens caerulescens) as they consumed underground stembases of tall cotton-grass (Eriophorum angustifolium) on an arctic staging area in northeastern Alaska. Geese fed in small patches ( =21.5 m²) of forage that made up 3% of the study area and consisted of high-quality aquatic graminoid and intermediate-quality wet sedge vegetation types. Dominant geese spent more time feeding in aquatic graminoid areas (r=0.61), but less total time feeding and more time resting than subdominant geese. Subdominant geese were displaced to areas of wet sedge where cotton-grass was a smaller proportion of underground biomass. Geese metabolized an average of 48% of the organic matter in stembases and there was a positive correlation between dominance and organic matter metabolizability (r=0.61). Total mean retention time of forage was 1.37 h and dry matter intake was 14.3 g/h. Snow geese that stage on the coastal plain of the Beaufort Sea likely use an extensive area because they consume a large mass of forage and exploit habitats that are patchily distributed and make up a small percentage of the landscape. Individual variation in nutrient absorption may result from agonistic interactions in an environment where resources are heterogeneously distributed.     

Breeding dispersal in a heterogeneous landscape: the influence of habitat and nesting success in greater snow geese

Despite numerous studies on breeding dispersal, it is still unclear how habitat heterogeneity and previous nesting success interact to determine nest-site fidelity at various spatial scales. In this context, we investigated factors affecting breeding dispersal in greater snow geese (Anser caerulescens atlanticus), an Arctic breeding species nesting in two contrasting habitats (wetlands and mesic tundra) with variable pattern of snowmelt at the time of settlement in spring. From 1994 to 2005, we monitored the nesting success and breeding dispersal of individually marked females. We found that snow geese showed a moderate amount of nest-site fidelity and considerable individual variability in dispersal distance over consecutive nesting attempts. This variability can be partly accounted for by the annual timing of snowmelt. Despite this environmental constraint, habitat differences at the colony level consistently affected nesting success and settlement patterns. Females nesting in wetlands had higher nesting success than those nesting in mesic tundra. Moreover, geese responded adaptively to spatial heterogeneity by showing fidelity to their nesting habitat, independently of snowmelt pattern. From year to year, geese were more likely to move from mesic to high-quality wetland habitat, regardless of previous nesting success and without cost on their subsequent nesting performance. The unpredictability of snowmelt and the low cost of changing site apparently favour breeding-site dispersal although habitat quality promotes fidelity at the scale of habitat patches.     

The role of sea ice for vascular plant dispersal in the Arctic

Sea ice has been suggested to be an important factor for dispersal of vascular plants in the Arctic. To assess its role for postglacial colonization in the North Atlantic region, we compiled data on the first Late Glacial to Holocene occurrence of vascular plant species in East Greenland, Iceland, the Faroe Islands and Svalbard. For each record, we reconstructed likely past dispersal events using data on species distributions and genetics. We compared these data to sea-ice reconstructions to evaluate the potential role of sea ice in these past colonization events and finally evaluated these results using a compilation of driftwood records as an independent source of evidence that sea ice can disperse biological material. Our results show that sea ice was, in general, more prevalent along the most likely dispersal routes at times of assumed first colonization than along other possible routes. Also, driftwood is frequently dispersed in regions that have sea ice today. Thus, sea ice may act as an important dispersal agent. Melting sea ice may hamper future dispersal of Arctic plants and thereby cause more genetic differentiation. It may also limit the northwards expansion of competing boreal species, and hence favour the persistence of Arctic species.     

Phylogeography of willow grouse (Lagopus lagopus) in the Arctic: taxonomic discordance as inferred from molecular data

Using independently segregating nuclear single nucleotide polymorphisms (SNPs) and mitochondrial control region sequences, we found an east–west division among sampled willow grouse Lagopus lagopus subspecies. This division cut across the range of the subspecies with the largest distribution (lagopus) and thus contradicted existing taxonomic classifications. Russian Lagopus lagopus lagopus tended to cluster with North American willow grouse partly classified as other subspecies. Scandinavian willow grouse (L. l. lagopus) clustered with red grouse from Britain and Ireland (Lagopus lagopus scoticus and Lagopus lagopus hibernicus) but substructuring confirmed the monophyly of the latter. In North America, we could not detect any major genetic divisions apart from two birds described as alexandrae from the Heceta Island (Alaska) when using mitochondrial sequences. Other samples from North America were intermingled regardless of whether they were described as muriei, alexandrae or lagopus. A specimen described as alexandrae was to some extent distinct when analysing the SNP data. The genetic analyses indicated some concordance between genetics and taxonomy but not complete congruence. This is particularly evident for mitochondrial DNA network analyses. We suggest that the taxonomy of this species would benefit by a careful re‐examination of the available evidence for subspecies. It appears as if subspecies status is a poor proxy for assigning evolutionary significant units and management units in this species. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 77–90.