Avian cholera in waterfowl: the role of lesser snow and ross's geese as disease carriers in the Playa Lakes Region

We collected samples from apparently healthy geese in the Playa Lakes Region (USA) during the winters of 2000-01 and 2001-02 to determine whether carriers of Pasteurella multocida, the bacterium that causes avian cholera, were present in wild populations. With the use of methods developed in laboratory challenge trials (Samuel et al., 2003a) and a serotype-specific polymerase chain reaction method for identification of P. multocida serotype 1, we found that a small proportion of 322 wild birds (<5%) were carriers of pathogenic P. multocida. On the basis of serology, an additional group of these birds (<10%) were survivors of recent avian cholera infection. Our results confirm the hypothesis that wild waterfowl are carriers of avian cholera and add support for the hypothesis that wild birds are a reservoir for this disease. In concert with other research, this work indicates that enzootic infection with avian cholera occurs in lesser snow goose (Chen caerulescens caerulescens) populations throughout their annual cycle. Although fewer Ross's geese (Chen rossii) were sampled, we also found these birds were carriers of P. multocida. Even in the absence of disease outbreaks, serologic evidence indicates that chronic disease transmission and recent infection are apparently occurring year-round in these highly gregarious birds and that a small portion of these populations are potential carriers with active infection.     

Of fleas and geese: the impact of an increasing nest ectoparasite on reproductive success

Past studies on the relationship between nest ectoparasites and avian fitness have been primarily limited to altricial hosts. Life history strategies of precocial and altricial birds vary considerably, limiting our ability to infer the effect of nest parasites on fitness of precocial species. Ross's Chen rossii and lesser snow goose Chen caerulescens caerulescens populations have been growing at unprecedented high rates. New limiting factors on vital rates of these precocial birds may arise after populations have been released from previously regulating factors. The flea Ceratophyllus vagabundus vagabundus is an apparently newly emerging nest parasite in the arctic goose colony at Karrak Lake, Nunavut, Canada. We examined the relationship between flea abundance (measured by the proportion of goose eggs covered by blood in each nest) and goose reproductive success from 2001–2004. In three of four years of study, nest success was inversely related to flea abundance in nests. Despite the potential for high costs to individuals, the overall effects of fleas on goose nesting success have thus far been small. We demonstrated that nest parasites negatively influence reproductive success of precocial bird hosts despite host life history strategy of leaving the nest quickly after hatch, which results in minimal exposure to nest parasites compared to altricial birds that raise their young in the nest.     

Antibodies against Pasteurella multocida in snow geese in the western Arctic.

To determine if lesser snow geese (Chen caerulescens caerulescens) are a potential reservoir for the Pasteurella multocida bacterium that causes avian cholera, serum samples and/or pharyngeal swabs were collected from > 3,400 adult geese breeding on Wrangel Island (Russia) and Banks Island (Canada) during 1993-1996. Pharyngeal swab sampling rarely (> 0.1%) detected birds that were exposed to P. multocida in these populations. Geese with serum antibody levels indicating recent infection with P. multocida were found at both breeding colonies. Prevalence of seropositive birds was 3.5% at Wrangel Island, an area that has no recorded history of avian cholera epizootics. Prevalence of seropositive birds was 2.8% at Banks Island in 1994, but increased to 8.2% during 1995 and 1996 when an estimated 40,000-60,000 snow geese were infected. Approximately 50% of the infected birds died during the epizootic and a portion of the surviving birds may have become carriers of the disease. This pattern of prevalence indicated that enzootic levels of infection with P. multocida occurred at both breeding colonies. When no avian cholera epizootics occurred (Wrangel Island, Banks Island in 1994), female snow geese (4.7%) had higher antibody prevalence than males (2.0%).     

Multi-species patterns of avian cholera mortality in Nebraska's Rainwater Basin

Nebraska's Rainwater Basin (RWB) is a key spring migration area for millions of waterfowl and other avian species. Avian cholera has been endemic in the RWB since the 1970s and in some years tens of thousands of waterfowl have died from the disease. We evaluated patterns of avian cholera mortality in waterfowl species using the RWB during the last quarter of the 20th century. Mortality patterns changed between the years before (1976-1988) and coincident with (1989-1999) the dramatic increases in lesser snow goose abundance and mortality. Lesser snow geese (Chen caerulescens caerulescens) have commonly been associated with mortality events in the RWB and are known to carry virulent strains of Pasteurella multocida, the agent causing avian cholera. Lesser snow geese appeared to be the species most affected by avian cholera during 1989-1999; however, mortality in several other waterfowl species was positively correlated with lesser snow goose mortality. Coincident with increased lesser snow goose mortality, spring avian cholera outbreaks were detected earlier and ended earlier compared to 1976-1988. Dense concentrations of lesser snow geese may facilitate intraspecific disease transmission through bird-to-bird contact and wetland contamination. Rates of interspecific avian cholera transmission within the waterfowl community, however, are difficult to determine.     

Evidence of Territoriality and Species Interactions from Spatial Point-Pattern Analyses of Subarctic-Nesting Geese

Quantifying spatial patterns of bird nests and nest fate provides insights into processes influencing a species’ distribution. At Cape Churchill, Manitoba, Canada, recent declines in breeding Eastern Prairie Population Canada geese (Branta canadensis interior) has coincided with increasing populations of nesting lesser snow geese (Chen caerulescens caerulescens) and Ross’s geese (Chen rossii). We conducted a spatial analysis of point patterns using Canada goose nest locations and nest fate, and lesser snow goose nest locations at two study areas in northern Manitoba with different densities and temporal durations of sympatric nesting Canada and lesser snow geese. Specifically, we assessed (1) whether Canada geese exhibited territoriality and at what scale and nest density; and (2) whether spatial patterns of Canada goose nest fate were associated with the density of nesting lesser snow geese as predicted by the protective-association hypothesis. Between 2001 and 2007, our data suggest that Canada geese were territorial at the scale of nearest neighbors, but were aggregated when considering overall density of conspecifics at slightly broader spatial scales. The spatial distribution of nest fates indicated that lesser snow goose nest proximity and density likely influence Canada goose nest fate. Our analyses of spatial point patterns suggested that continued changes in the distribution and abundance of breeding lesser snow geese on the Hudson Bay Lowlands may have impacts on the reproductive performance of Canada geese, and subsequently the spatial distribution of Canada goose nests.     

Foraging time and dietary intake by breeding Ross's and Lesser Snow Geese

We compared foraging times of female Ross's (Chen rossii) and Lesser Snow Geese (Chen caerulescens caerulescens) breeding at Karrak Lake, NT, Canada and examined variation due to time of day and reproductive stage. We subsequently collected female geese that had foraged for known duration and we estimated mass of foods consumed during foraging bouts. Female Ross's Geese spent more time foraging (mean % - SE =28.4ǃ.3%; P=0.0002), on average, than did female Lesser Snow Geese (21.5 - 1.4%). Foraging time by female geese differed among reproductive stages, but differences were not consistent among time periods (stage-by-time block interaction, P=0.0003). Females spent considerably more time foraging during prelaying and laying than during incubation. Ross's Geese also spent a greater percent of time feeding (83.0DŽ.8%) during incubation recesses than did Lesser Snow Geese (60.9Dž.6%). Consumption of organic matter during foraging bouts was minimal; estimated consumption averaged 9.6dž.0 and 12.4dž.6 g (mean - SE) dry mass/day before incubation and 5.9DŽ.0 and 5.7DŽ.1 g dry mass/day during incubation for Lesser Snow and Ross's Geese, respectively. Diets consisted primarily of mosses (bryophytes), Chickweed (Stellaria spp.) and Sedges (Carex spp.). Before incubation, eggshell consumption was estimated as 4.3Dž.2 and 0.4ǂ.3 g dry mass/day for Lesser Snow and Ross's Geese, respectively; neither species consumed eggshell during incubation. We conclude that eggshell from nests of previous years is likely an important source of dietary calcium used to meet mineral demands of eggshell formation at Karrak Lake. Our findings of wide disparities between foraging time and food intake indicate that results from studies that do not directly measure intake rates remain equivocal. Finally, we propose four hypotheses accounting for foraging effort that evidently yields little nutritional or energetic benefit to geese nesting at Karrak Lake.     

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.     

Does body size influence nest attendance? A comparison of Ross’s geese (Chen rossii) and the larger, sympatric lesser snow geese (C. caerulescens caerulescens)

The body-size hypothesis predicts that nest attendance is positively related to body size among waterfowl and that recess duration is inversely related to body size. Several physiological and behavioral characteristics of Ross’s geese (Chen rossii) suggest that females of this species should maintain high nest attendance despite their relatively small body size. Accordingly, we used 8-mm films to compare the incubation behavior of Ross’s geese to that of the larger, closely-related lesser snow geese (C. caerulescens caerulescens; hereafter, snow geese) nesting sympatrically at Karrak lake, Nunavut, Canada in 1993. We found that nest attendance averaged 99% for both species. Our results offer no support for the body-size hypothesis. We suggest that temperature requirements of embryos in relation to short incubation duration and a low foraging efficiency of females select for high nest attendance in both snow geese and Ross’s geese.     

The genetic legacy of Mother Goose– phylogeographic patterns of lesser snow goose Chen caerulescens caerulescens maternal lineages

By using the polymerase chain reaction to amplify and sequence 178 bp of a rapidly evolving region of the mtDNA genome (segment I of the control region) from 81 individuals, approximately 11% of the variation present in the lesser snow goose Chen caerulescens caerulescens L. mitochondrial genome was surveyed. The 26 types of mtDNA detected formed two distinct mitochondrial clades that differ by an average of 6.7% and are distributed across the species range. Restriction analysis of amplified fragments was then used to assign the mtDNA of an additional 29 individuals to either of these clades. Within one major clade, sequence among mtDNAs was concordant with geographic location. Within the other major clade the degree of sequence divergence among haplotypes was lower and no consistent geographic structuring was evident. The two major clades presumably result from vicariant separation of lesser snow geese during the Pleistocene.     

Do geese fully develop brood patches? A histological analysis of lesser snow geese (<Emphasis Type="Italic">Chen caerulescens caerulescens</Emphasis>) and Ross’s geese (<Emphasis Type="Italic">C. rossii</Emphasis>)

Most birds develop brood patches before incubation; epidermis and dermis in the brood patch region thicken, and the dermal connective tissue becomes increasingly vascularized and infiltrated by leukocytes. However, current dogma states that waterfowl incubate without modifications of skin within the brood patch region. The incubation periods of lesser snow geese (Chen caerulescens caerulescens; hereafter called snow geese) and Ross’s geese (C. rossii) are 2–6 days shorter than those of other goose species; only females incubate. Thus, we hypothesized that such short incubation periods would require fully developed brood patches for sufficient heat transfer from incubating parents to eggs. We tested this hypothesis by analyzing the skin histology of abdominal regions of snow and Ross’s geese collected at Karrak Lake, Nunavut, Canada. For female snow geese, we found that epidermis and dermis had thickened and vascularization of dermis was 14 times greater, on average, than that observed in males (n=5 pairs). Our results for Ross’s geese (n=5 pairs) were more variable, wherein only one of five female Ross’s geese fully developed a brood patch. Our results are consistent with three hypotheses about brood patch development and its relationship with different energetic cost–benefit relationships, resulting from differences in embryonic development and body size.     

Impact of migratory snow geese on nitrogen and phosphorus dynamics in a freshwater reservoir

1. We examined impacts of nutrient loading, particularly of nitrogen and phosphorus, from greater snow geese (Chen caerulescens atlantica) on a reservoir in south‐eastern Pennsylvania, U.S.A. Approximately 100 000 geese use the reservoir for 2–6 weeks prior to their spring migration northward. 2. We estimated the magnitude of nutrient loading by geese during their presence and compared that to surface input and output rates. We also conducted nutrient limitation bioassay experiments to examine patterns of algal nutrient limitation upstream and downstream of the reservoir. 3. During their presence from 1 February to 27 March 2001, snow geese contributed 85–93% of the phosphorus and 33–44% of the nitrogen loaded to the reservoir. Both nutrients were exported from the reservoir slowly rather than as a quick pulse. Consequently, phosphorus concentrations in the outflow were higher than in the inflow from February to the end of July. However, nitrogen concentrations were consistently lower in the outflow than the inflow. 4. Nutrient limitation bioassays conducted in June and July indicated that primary production in the outflow was limited by nitrogen whereas the inflow was co‐limited by nitrogen and phosphorus. Further downstream from the reservoir, primary production was consistently phosphorus limited. Therefore, nitrogen limitation persisted long after the geese had left, but was relatively localised.     

Associations between body composition and helminths of lesser snow geese during winter and spring migration

Costs of parasitism are predicted to be higher with greater parasite intensities and higher inter-parasite competition (diversity). We tested whether greater helminth intensities and diversity were associated with poorer body composition (whole-body fat, protein, mineral and true body mass) in lesser snow geese, Chen caerulescens caerulescens. As part of a larger study on nutritional ecology, 828 wintering or migrating geese were shot between January and May 1983 in 27 different date-locations (samples) during their northward migration through mid-continental North America. A large proportion of overall variation in body composition and parasite communities was among samples, so we analyzed data within each of the 27 samples, controlling for structural body size (the first principal component of 10 body size measurements), sex and the age of geese. There was no compelling evidence that cestodes, trematodes or helminth diversity were associated with variation in body composition but nematodes had several negative associations with fat reserves. However, negative associations between fat reserves and nematodes occurred most often in geese collected between March and May when nematode prevalences and intensities were relatively low. This suggests several possibilities: that the most common nematodes (Heterakis dispar and Trichostrongylus tenuis) were more virulent at this time, that infected individuals had been chronically infected and suffered cumulative nutrient deficits that lasted until late in the spring migration, or that geese became more vulnerable to the effects of parasites at this time of year, possibly because they redirected resources away from immunity toward fat storage in preparation for reproduction.     

Time budgets of Snow Geese Chen caerulescens and Ross's Geese Chen rossii in mixed flocks: implications of body size, ambient temperature and family associations

Body size affects foraging and forage intake rates directly via energetic processes and indirectly through interactions with social status and social behaviour. Ambient temperature has a relatively greater effect on the energetics of smaller species, which also generally are more vulnerable to predator attacks than are larger species. We examined variability in an index of intake rates and an index of alertness in Lesser Snow Geese Chen caerulescens caerulescens and Ross's Geese Chen rossii wintering in southwest Louisiana. Specifically we examined variation in these response variables that could be attributed to species, age, family size and ambient temperature. We hypothesized that the smaller Ross's Geese would spend relatively more time feeding, exhibit relatively higher peck rates, spend more time alert or raise their heads up from feeding more frequently, and would respond to declining temperatures by increasing their proportion of time spent feeding. As predicted, we found that Ross's Geese spent more time feeding than did Snow Geese and had slightly higher peck rates than Snow Geese in one of two winters. Ross's Geese spent more time alert than did Snow Geese in one winter, but alert rates differed by family size, independent of species, in contrast to our prediction. In one winter, time spent foraging and walking was inversely related to average daily temperature, but both varied independently of species. Effects of age and family size on time budgets were generally independent of species and in accordance with previous studies. We conclude that body size is a key variable influencing time spent feeding in Ross's Geese, which may require a high time spent feeding at the expense of other activities.     

A general model for the analysis of mark-resight, mark-recapture, and band-recovery data under tag loss

Estimates of waterfowl demographic parameters often come from resighting studies where birds fit with individually identifiable neck collars are resighted at a distance. Concerns have been raised about the effects of collar loss on parameter estimates, and the reliability of extrapolating from collared individuals to the population. Models previously proposed to account for collar loss do not allow survival or harvest parameters to depend on neck collar presence or absence. Also, few models have incorporated recent advances in mark-recapture theory that allow for multiple states or auxiliary encounters such as band recoveries. We propose a multistate model for tag loss in which the presence or absence of a collar is considered as a state variable. In this framework, demographic parameters are corrected for tag loss and questions related to collar effects on survival and recovery rates can be addressed. Encounters of individuals between closed sampling periods also can be incorporated in the analysis. We discuss data requirements for answering questions related to tag loss and sampling designs that lend themselves to this purpose. We illustrate the application of our model using a study of lesser snow geese (Chen caerulescens caerulescens).     

Mortality of geese as a result of collision with the ground

Two incidents are reported in which groups of migrating wild geese were found dead in agricultural fields in southern Manitoba during spring. In each case, the birds died overnight and poisoning was suspected; however, the birds had lesions of severe traumatic injury. The first incident, in 1985, involved about 150 lesser snow geese (Anser caerulescens caerulescens); the second, in 2003, involved 62 Canada geese (Branta canadensis). Both incidents occurred on dark, moonless nights. One possible explanation is that the birds became disoriented in a manner analogous to spatial disorientation described in aircraft pilots and flew as a flock directly into the earth. In the first incident, geese might have been frightened by sonic booms from aircraft; in the second, there was a thunderstorm with strong gusty winds in the area.     

New species records and changes in abundance of waterfowl in northwest Greenland

Breeding populations of Nearctic and Palearctic waterfowl have undergone significant changes in abundance and distribution over the past 50 years. The Avanersuaq District in northwest Greenland is home to an assemblage of waterfowl from both geographic areas; however, minimal historic or current information is available on species abundance. In 2008 and 2009, we conducted field surveys in Greenland from 76.00° to 77.35°N for breeding and non-breeding waterfowl and have collected anecdotal field notes of avian observations over a 20-year period (1993–2012). During these periods, we documented the first observation of a Ross’s goose (Chen rossii) and the first confirmed breeding by lesser snow geese (Chen caerulescens caerulescens) in Greenland. Northern pintails (Anas acuta) were observed for the first time in northwest Greenland, and a previously unknown breeding location for brent geese (Branta bernicla hrota) was also identified. Local populations of greater snow (C. c.) and Canada geese (B. canadensis) have increased in size. The Booth Sound and Drown Bay wetland areas and many islands throughout the Avanersuaq District were identified as critical habitat for both breeding and non-breeding waterfowl. Further increases in waterfowl abundance, including more frequent rare and new visitors, are likely in the study area as breeding populations further south continue to increase and an ameliorating climate allows for a longer breeding season. These results will prove useful as a baseline for comparisons with future surveys.     

Trophic interactions in a high arctic snow goose colony

We examined the role of trophic interactions in structuringa high arctic tundra community characterized by a large breedingcolony of greater snow geese (Chen caerulescens atlantica).According to the exploitation ecosystem hypothesis of Oksanenet al. (1981), food chains are controlled by top-down interactions.However, because the arctic primary productivity is low, herbivorepopulations are too small to support functional predator populationsand these communities should thus be dominated by the plant/herbivore trophic-level interaction. Since 1990, we have beenmonitoring annual abundance and productivity of geese, the impactof goose grazing, predator abundance (mostly arctic foxes, Alopexlagopus) and the abundance of lemmings, the other significantherbivore in this community, on Bylot Island, Nunavut, Canada.Goose grazing consistently removed a significant proportionof the standing crop (     

Do foraging methods in winter affect morphology during growth in juvenile snow geese?

Physical exertion during growth can affect ultimate size and density of skeletal structures. Such changes from different exercise regimes may explain morphological differences between groups, such as those exhibited by lesser snow geese (Chen caerulescens caerulescens; hereafter snow geese) foraging in southwest Louisiana. In rice‐prairie habitats (hereafter rice‐prairies), snow geese bite off or graze aboveground vegetation, whereas they dig or grub for subterranean plant parts in adjacent coastal marshes. Grubbing involves considerably more muscular exertion than does grazing. Thus, we hypothesized that rates of bone formation and growth would be lower for juveniles wintering in rice‐prairies than those in coastal marshes, resulting in smaller bill and skull features at adulthood. First, we tested this exertion hypothesis by measuring bills, skulls, and associated musculature from arrival to departure (November–February) in both habitats in southwest Louisiana, using both banded birds and collected specimens. Second, we used the morphological data to test an alternative hypothesis, which states that smaller bill dimensions in rice‐prairies evolved because of hybridization with Ross's geese (C. rossii). Under the exertion hypothesis, we predicted that bill and skull bones of juveniles would grow at different rates between habitats. However, we found that bill and skull bones of juveniles grew similarly between habitats, thus failing to support the exertion hypothesis. Morphometrics were more likely to differ by sex or change with sampling date than to differ by habitat. We predicted that significant, consistent skewness toward smaller birds could indicate hybridization with Ross's geese, but no skewness was observed in our morphological data, which fails to support the hybridization hypothesis. Further research is needed to clarify whether snow geese wintering in Louisiana represent a single polymorphic population that segregates into individually preferred habitats, which we believe at present to be more likely as an explanation than two ecologically and spatially distinct morphotypes.     

Correlations between sequential timing decisions do not necessarily indicate strategic behavior: a comment on Bêty et al

Using correlation analysis, Bêty, Gauthier, and Giroux confirmed that individuals of the migrating snow geese (Chen caerulescens atlantica) arriving earlier on their breeding grounds experienced longer prelaying periods but laid earlier than late-arriving birds. They concluded that these birds strategically adjusted timing to optimize fitness. This test is, however, not sufficient to indicate strategic behavior. We show analytically that, without assuming strategic behavior, arrival date, prelaying period, and laying date are correlated with each other because of their sequential occurrence. We conclude that correlation tests on sequentially occurring variables are not sufficient to detect nonrandom relations.