Occupancy time in sets of states for demographic models.

As an individual moves through its life cycle, it passes through a series of states (age classes, size classes, reproductive states, spatial locations, health statuses, etc.) before its eventual death. The occupancy time in a state is the time spent in that state over the individual's life. Depending on the life cycle description, the occupancy times describe different demographic variables, for example, lifetime breeding success, lifetime habitat utilisation, or healthy longevity. Models based on absorbing Markov chains provide a powerful framework for the analysis of occupancy times. Current theory, however, can completely analyse only the occupancy of single states, although the occupancy time in a set of states is often desired. For example, a range of sizes in a size-classified model, an age class in an age×stage model, and a group of locations in a spatial stage model are all sets of states. We present a new mathematical approach to absorbing Markov chains that extends the analysis of life histories by providing a comprehensive theory for the occupancy of arbitrary sets of states, and for other demographic variables related to these sets (e.g., reaching time, return time). We apply this approach to a matrix population model of the Southern Fulmar (Fulmarus glacialoides). The analysis of this model provides interesting insight into the lifetime number of breeding attempts of this species. Our new approach to absorbing Markov chains, and its implementation in matrix oriented software, makes the analysis of occupancy times more accessible to population ecologists, and directly applicable to any matrix population models.

Impact of Age, Season, and Flowing vs. Stagnant Water Habitat on Avian Influenza Prevalence in Mute Swan (Cygnus olor) in Belgium.

Due to their probable role in the spread of Asian highly pathogenic avian influenza (HPAI) H5N1 virus, and in order to explore its implication in the low pathogenic avian influenza (LPAI) virus epidemiology, mute swans represent one particular wild bird species specifically targeted in the avian influenza (AI) surveillance elaborated in Belgium. A total of 640 individual mute swans have been sampled during a 4-yr AI surveillance program (2007-2010) to determine the AI seroprevalence and viroprevalence in this species; all were analyzed through age, temporal, and habitat (flowing and stagnant water) factors. Using a nucleoprotein (NP)-based ELISA, a global antibody prevalence of 35% has been found and was characterized by two peaks in the winter and the summer that might be indicative of a greater LPAI virus circulation in the autumn than in the spring. A significantly higher antibody prevalence was detected in adult swans (53.8%) as compared to juveniles (15.5%). In contrast, a low prevalence of infection (2.7%) was found, mainly in juvenile mute swans and only during the autumn migration period. Interestingly, an impact of water habitat was observed based on the comparison of the antibody prevalence and prevalence of infection from swan populations living on stagnant water vs. flowing water, suggesting that stagnant water provides a more-favorable environment for LPAI persistence and transmission.

Widespread exposure to lead affects the body condition of free-living whooper swans Cygnus cygnus wintering in Britain.

Lead poisoning, through the ingestion of spent lead gunshot, is an established cause of morbidity and mortality in waterbirds globally, but the thresholds at which blood levels begin to affect the physiology of birds in the wild are less well known. Here we determine the prevalence of lead exposure in whooper swans and, for the first time, identify the level of blood lead associated with initial reductions in body condition. Blood lead elevated above background levels (i.e. >20 µg dL(-1)) was found in 41.7% (125/300) of swans tested. Blood lead was significantly negatively associated with winter body condition when levels were ≥44 µg dL(-1) (27/260 = 10%). Our findings indicating that sub-lethal impacts of lead on body condition occur at the lower end of previously established clinical thresholds and that a relatively high proportion of individuals in this population may be affected, reaffirm the importance of reducing contamination of the environment with lead shot.

Improving residual feed intake of mule progeny of Muscovy ducks: genetic parameters and responses to selection with emphasis on carcass composition and fatty liver quality.

In France, about 95% of the fatty liver production comes from mule ducks, an infertile hybrid cross between female common ducks (Anas platyrhynchos) and Muscovy drakes (Cairina moschata). As a hybrid, genetic improvement of the performance of mule ducks is achieved by selection of the parental populations. Feed represents two-thirds of the total costs of duck production; thus, the breeders' main concern is to improve the birds' feed efficiency without impairing their capacity to be overfed after the growing period. To obtain insight into this economically important trait, we designed a divergent selection experiment for Muscovy sires on the basis of the residual feed intake (RFI) of their male mule progeny. Residual feed intake represents the fraction of feed intake that cannot be explained by maintenance and production requirements. Data comprised records from generations 0 to 2 with 227 Muscovy sires of 2,016 mule ducks. Growth and fat deposition were measured individually for all mule ducks. Half-sib mule ducks were placed together in 1 small pen, and the feed consumption was recorded for each pen; all half-sibs therefore had the same feed intake estimates. Heritability of RFI was high (0.83 ± 0.42), and after 2 generations of selection, the differences between lines in terms of RFI (contrast of 9.13 g/d, representing 0.76 phenotypic standard deviation; P < 0.0001) and feed conversion ratio (contrast of 0.08, representing 0.28 phenotypic standard deviation; P = 0.0003) were marked. In regard to carcass traits, the low-RFI line (high-efficiency animals) had heavier legs (476 vs. 463 g; P < 0.0001) and magret muscles (268 vs. 262 g; P = 0.001) than the high-RFI line (low-efficiency animals). No differences were observed for either the liver weight or the melting rate between the 2 lines (P > 0.46). Selection on RFI thus improves the feed efficiency of animals without impairing their capacity to produce fatty liver.

Abiotic factors affecting the persistence of avian influenza virus in surface waters of waterfowl habitats.

Avian influenza (AI) virus can remain infectious in water for months, and virus-contaminated surface water is considered to be a source of infection within wild waterfowl populations. Previous work has characterized the effects of pH, salinity, and temperature on viral persistence in water, but most of that work was done with modified distilled water. The objective of this study was to identify the abiotic factors that influence the duration of AI virus persistence in natural surface water. Surface water samples were collected from 38 waterfowl habitats distributed across the United States. Samples were submitted to the U.S. Geological Survey National Water Quality Laboratory for chemical analysis and the University of Georgia for viral reduction time analysis. Samples were filtered with 0.22-µm filters, and the durations of persistence of three wild-bird-derived influenza A viruses within each water sample at 10, 17, and 28°C were determined. The effects of the surface water physicochemical factors on the duration of AI viral persistence in laboratory experiments were evaluated by multivariable linear regression with robust standard errors. The duration of AI virus persistence was determined to be longest in filtered surface water with a low temperature (<17°C), a neutral-to-basic pH (7.0 to 8.5), low salinity (<0.5 ppt), and a low ammonia concentration (<0.5 mg/liter). Our results also highlighted potential strain-related variation in the stability of AI virus in surface water. These results bring us closer to being able to predict the duration of AI virus persistence in surface water of waterfowl habitats.

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. 

Histopathology of liver and kidneys of wild living Mallards Anas platyrhynchos and Coots Fulica atra with considerable concentrations of lead and cadmium.

Concentrations of cadmium and lead were measured in liver and kidneys of Mallard (n=60) and Coot (n=50). Free living birds were collected by hunters in years 2006-2008 in the area of fishponds near Zator in southern Poland. Age group was determined according to the appearance of the plumage (Mallards) and iris color (Coot). Concentrations of metals were measured with ET-AA spectrometer. Among all birds specimens with negligible (n=5) and high concentrations (Mallards n=18 and Coots n=17) of cadmium and lead were chosen for further analysis. Histopathological alterations were observed, ranging from circulatory disturbances, retrogressive changes, inflammations to leukocytic infiltration in liver and kidney. They dominated among birds with the highest concentrations of metals. The control group of birds was characterized by a very small number of mentioned lesions. Probably the higher cadmium and lead concentrations in tissues are co-factors in the development of lesions.

Influence of pasture intake on meat quality, lipid oxidation, and fatty acid composition of geese.

This study was conducted to investigate the influence of pasture intake on meat quality, lipid oxidation, and fatty acid composition of geese. One hundred twenty Dongbei White male geese (a local breed; BW = 878 ± 13 g; 28 d old) were randomly and equally divided into 2 treatments with 6 pens of 10 geese per treatment. The 2 treatments consisted of birds fed ad libitum a corn-based feed. One-half of the birds had no access to pasture (control) while the other half had access to an alfalfa (Medicago sativa)-based pasture (pasture). The study lasted 42 d. Body weight and feed intake were recorded weekly. At the end of the study, geese were slaughtered to collect meat samples. Results showed that pasture intake reduced subcutaneous fat thickness (P < 0.05) and abdominal fat yield (P < 0.05) of geese compared with control. Geese with access to pasture had greater cooking loss (P < 0.05) and lightness (L*) value (P < 0.05) and lower pH at 24 h postmortem (pH24; P < 0.05) and thiobarbituric acid reacting substance values (P < 0.05) at 0 and 30 min of forced oxidation. Moreover, pasture intake increased linolenic acid (C18:3n-3; P < 0.05) and eicosapentaenoic acid (C20:5n-3; P < 0.05) and reduced the n-6:n-3 ratio (P < 0.05) in the breast muscle of geese compare with the control. In conclusion, pasture intake did not enhance growth performance but improved carcass characteristics and meat quality and changed fatty acid composition of geese.

Preliminary studies on the reaction of growing geese (Anser anser f. domestica) to the proximity of wind turbines.

Wind farms produce electricity without causing air pollution and environmental degradation. Unfortunately, wind turbines are a source of infrasound, which may cause a number of physiological effects, such as an increase in cortisol and catecholamine secretion. The impact of infrasound noise, emitted by wind turbines, on the health of geese and other farm animals has not previously been evaluated. Therefore, the aim of this study was to determine the effect of noise, generated by wind turbines, on the stress parameters (cortisol) and the weight gain of geese kept in surrounding areas. The study consisted of 40 individuals of 5-week-old domestic geese Anser anser f domestica, divided into 2 equal groups. The first experimental gaggle (I) remained within 50 m from turbine and the second one (II) within 500 m. During the 12 weeks of the study, noise measurements were also taken. Weight gain and the concentration of cortisol in blood were assessed and significant differences in both cases were found. Geese from gaggle I gained less weight and had a higher concentration of cortisol in blood, compared to individuals from gaggle II. Lower activity and some disturbing changes in behavior of animals from group I were noted. Results of the study suggest a negative effect of the immediate vicinity of a wind turbine on the stress parameters of geese and their productivity.

Effects of presentation and type of cereals (corn or sorghum) on performance of geese.

The aim of this trial was to study the effects of cereal (corn or sorghum) and of the presentation of the diet (pelleted diet or a mixture with cereal) on the performance of geese. In total, 480 one-day-old geese were divided into 4 groups differing in the diet they received between 52 and 102 d of age: a pelleted diet containing 500 g of sorghum/kg (CS group; AMEn 11.29 MJ/kg, CP 16.70%); a mixture containing 500 g of protein-rich pellets and 500 g of sorghum whole seeds/kg (MS group; AMEn 11.61 MJ/kg, CP 14.30%); a pelleted diet containing 500 g of corn/kg (group CC; AMEn 11.33 MJ/kg, CP 16.40%); and a mixture containing 500 g of protein-rich pellets and 500 g of corn mash/kg (group MC; AMEn 11.48 MJ/kg, CP 14.50%). From 102 to 117 d, 33 birds/sex/group were force-fed with corn. Between 52 and 102 d of age, individual BW and collective feed intake (40 birds/pen) were measured weekly. Body traits were measured at 69, 96, and 117 d and the volume of the crop was measured at 102 d of age. The cereal had no significant effect on total feed intake (15,028 g, NS) or BW (5,811 g, NS) but the breast development at 102 d was less in birds fed sorghum compared with corn (-4.9%; P<0.05). Feed intake (+8.4% between 69 and 102 d; P<0.001), BW (+3.9%; P<0.001) and gut development (+9.3%; P<0.001) were higher when the diet was offered as a mixture compared with a complete pelleted diet. The birds fed the complete pelleted diets had a larger crop (+10.1%; P<0.05) than others. The cereal and the presentation of the diet had no effect on the body traits at the end of the force feeding. The results suggest that the use of sorghum whole seeds is interesting for feeding geese.

[Community composition, seasonal dynamics and interspecific correlation of waterbirds in the Qiantangjiang River estuary and Hangzhou Bay].

Waterbird surveys were conducted regularly in the Qiantangjiang River estuary and Hangzhou Bay from July 2007 to November 2011. A total of 128 species (nine orders and 18 families) were recorded, including 119 migrants which accounted for 93% of the total species; eleven species were listed as National Protected Species. Inter-specific correlation analysis for 13 shorebird populations and nine duck populations recorded over time found that 21 pairs of shorebirds and 23 pairs of ducks were correlated. By looking at seasonal dynamics and migration patterns we were able to divide the migration process into six stages: (1) late July to late September was the migration peak of shorebirds, which were dominated by Limosa limosa, Calidris ruficollis and Charadrius mongolus. (2) Early October to mid-December was the migration peak of wintering migrants of shorebirds and ducks, which were the first two large groups in our study areas. (3) Late December to mid-February was the wintering period of migration waterbirds. (4) Late February to late March was the peak migration of ducks and the winter migrants of shorebirds dominated by Calidris alpina. (5) Early April to mid-May was the migration peak of passage migrants such as, Calidris ruficollis, Calidris acuminate and Limosa limosa but the population size of shorebird winter migrants dominated by Calidris alpine was still larger than the former. (6) Late May to mid-July was the breeding season of all egrets, summer migrants of gulls and several species of shorebirds. Our surveys show that interaction among species is possibly an important determinant of community composition of shorebirds and wintering ducks during the migration season. It may be the geographical position and community composition of migrant shorebirds across Hangzhou Bay that mean during the northward migration there are far more shorebirds than during the southward migration.

[Waterbird habitat-selection during winter and spring in reclaimed coastal wetlands in Nanhui Dongtan, Shanghai].

From December 2009 to May 2010 goose and duck (Anatidae) community censuses in winter and shorebird (Charadriiforms) community censuses in spring were conducted across three types artificial wetlands (urban lake wetland, restorative wetland, abandoned wetland) along the coast of Nanhui, Shanghai. Correlation analyses were undertaken between community indices and habitat factors. The results showed there were significant differences in the density of geese and ducks among the wetlands, but no difference in the number of species. The density of geese and ducks in the restorative wetland was 3.77 times that of abandoned wetland and 6.03 times that of urban lake wetlands. The number of species and density of shorebirds in restorative wetlands was 2.88 and 5.70 times that of abandoned wetlands. We found significant differences in the number and density of shorebird species between restorative and abandoned wetlands. The number of species density of geese and ducks and the Shannon-Wiener (H') index were positively correlated with water area. The number of species and H' were negatively correlated with vegetation area. The number of species, species density and H' and evenness were negatively correlated with vegetation coverage. H' was positively correlated with mean water level. The results showed that the number and density of shorebird species were positively correlated with bare muddy areas. Aquaculture ponds and paddy fields in reclaimed area is efficient sufficient compensation mechanism to maintain more water areas for waterbirds and to control vegetation expansion and maintain shorebird habitat after coastal reclamation.

Mercury demethylation in waterbird livers: dose-response thresholds and differences among species.

We assessed methylmercury (MeHg) demethylation in the livers of adults and chicks of four waterbird species that commonly breed in San Francisco Bay: American avocets, black-necked stilts, Caspian terns, and Forster's terns. In adults (all species combined), we found strong evidence for a threshold model where MeHg demethylation occurred above a hepatic total mercury concentration threshold of 8.51 +/- 0.93 microg/g dry weight, and there was a strong decline in %MeHg values as total mercury (THg) concentrations increased above 8.51 microg/g dry weight. Conversely, there was no evidence for a demethylation threshold in chicks, and we found that %MeHg values declined linearly with increasing THg concentrations. For adults, we also found taxonomic differences in the demethylation responses, with avocets and stilts showing a higher demethylation rate than that of terns when concentrations exceeded the threshold, whereas terns had a lower demethylation threshold (7.48 +/- 1.48 microg/g dry wt) than that of avocets and stilts (9.91 +/- 1.29 microg/g dry wt). Finally, we assessed the role of selenium (Se) in the demethylation process. Selenium concentrations were positively correlated with inorganic Hg in livers of birds above the demethylation threshold but not below. This suggests that Se may act as a binding site for demethylated Hg and may reduce the potential for secondary toxicity. Our findings indicate that waterbirds demethylate mercury in their livers if exposure exceeds a threshold value and suggest that taxonomic differences in demethylation ability may be an important factor in evaluating species-specific risk to MeHg exposure. Further, we provide strong evidence for a threshold of approximately 8.5 microg/g dry weight of THg in the liver where demethylation is initiated.

Paternity assessment: application on estimation of breeding value in body-weight at first egg trait of egg-laying duck (Anas platyrhynchos).

Paternity index was analyzed using five microsatellite loci among Chinese egg-laying ducks (Anas platyrhynchos). Based on the paternity relationship that was identified by paternity index analysis, the estimated breeding value (EBV) was calculated using BLUP (best linear unbiased predictor) method. Body weight at first egg (BWF) is the only considered trait in this study. In total, 12 sires, 31 dams and 77 daughters were involved in the EBV calculation. The results demonstrated that five microsatellite loci's polymorphism information content (PIC) ranged from 0.795 in locus AY493338 to 0.957 in locus AY493264 with average 0.899; the parent-offspring relationships were built by these microsatellites' genotype, 12 families of half sibling and 2 families of full sibling were involved, and the relationship error is smaller than 10(-7). The EBV results suggest that the average EBV was significantly higher in females (average EBV is 10.234 and 0.1045 for mother and daughter, respectively) than males (average EBV is just -26.44). The EBV results on BWF were in good agreement with the principle of GH (growth hormone) expression in poultry. These results show that paternity analyses of Chinese egg-laying ducks were basically resolved using the five microsatellite loci selected. The paternity relationships can apply in Chinese egg-laying duck breeding to quicken the improvement of genetic progress.

Vitamin A and contaminant concentrations in surf scoters (Melanitta perspicillata) wintering on the Pacific coast of British Columbia, Canada.

Surf scoters are part of a community of sea ducks on the western coast of North America that have shown signs of long-term, unexplained declines in breeding bird numbers. Substantial numbers of scoters winter in the major harbours on the west coast, after breeding in the west-central northern boreal forest. To address the potential for contaminants to impact the health and survival of those birds, we investigated the condition and contamination of surf scoters during the winters of 1998-2001 at four foraging locations in the Strait of Georgia region of the Pacific coast of Canada. Vitamin A status was evaluated in liver and plasma samples collected from adults and juveniles, as part of a larger assessment of tissue contamination, body condition and biomarker responses. Individuals collected from a relatively contaminated site, Howe Sound, showed consistently low hepatic concentrations of retinol and retinyl palmitate forms of vitamin A, and gender-specific associations of retinyl palmitate with hepatic EROD activity. The relationship of hepatic retinol to retinyl palmitate was not constant across geographic locations, and a clear, linear relationship between the two forms of vitamin A was only evident in birds from the relatively uncontaminated site. This study also identified strong positive relationships between vitamin A and tissue burdens of cadmium and zinc. The positive association between hepatic retinyl palmitate and renal cadmium is similar to one observed in laboratory rats, in which a mechanism of interference with the controlled release of retinol from the liver was suggested.

Evolutionary response to selection on clutch size in a long-term study of the mute swan.

Life-history traits in wild populations are often regarded as being subject to directional selection, and the existence of substantial variation and microevolutionary stasis of these characters is therefore a problem in need of explanation. Avian clutch size is an archetypal life-history trait in this context, and many studies have sought to test explanations for stasis in clutch size. Surprisingly, there are many fewer studies that used long-term data to ask how selection acts on clutch size, particularly in a multivariate framework. In this article, we report selection, inheritance, and evolution of clutch size over 25 years in a colony of mute swans using a multivariate quantitative genetic framework to control for correlations with breeding time. We show that clutch size is influenced by both additive genetic and permanent environmental effects and that selection acts on clutch size in combination with breeding time. Natural selection on clutch size is strongly directional, favoring larger clutches, and we observe an increase in clutch size of 0.35 standard deviations, consistent with the expected response based on selection and inheritance of clutch size. We hypothesize that these changes result from recent relaxation of food constraints and predation risks experienced by this colony.

Quantitative genetics of age at reproduction in wild swans: support for antagonistic pleiotropy models of senescence.

Why do individuals stop reproducing after a certain age, and how is this age determined? The antagonistic pleiotropy theory for the evolution of senescence predicts that increased early-life performance should be accompanied by earlier (or faster) senescence. Hence, an individual that has started to breed early should also lose its reproductive capacities early. We investigate here the relationship between age at first reproduction (AFR) and age at last reproduction (ALR) in a free-ranging mute swan (Cygnus olor) population monitored for 36 years. Using multivariate analyses on the longitudinal data, we show that both traits are strongly selected in opposite directions. Analysis of the phenotypic covariance between these characters shows that individuals vary in their inherent quality, such that some individuals have earlier AFR and later ALR than expected. Quantitative genetic pedigree analyses show that both traits possess additive genetic variance but also that AFR and ALR are positively genetically correlated. Hence, although both traits display heritable variation and are under opposing directional selection, their evolution is constrained by a strong evolutionary tradeoff. These results are consistent with the theory that increased early-life performance comes with faster senescence because of genetic tradeoffs.

Age-dependent genetic variance in a life-history trait in the mute swan.

Genetic variance in characters under natural selection in natural populations determines the way those populations respond to that selection. Whether populations show temporal and/or spatial constancy in patterns of genetic variance and covariance is regularly considered, as this will determine whether selection responses are constant over space and time. Much less often considered is whether characters show differing amounts of genetic variance over the life-history of individuals. Such age-specific variation, if present, has important potential consequences for the force of natural selection and for understanding the causes of variation in quantitative characters. Using data from a long-term study of the mute swan Cygnus olor, we report the partitioning of phenotypic variance in timing of breeding (subject to strong natural selection) into component parts over 12 different age classes. We show that the additive genetic variance and heritability of this trait are strongly age-dependent, with higher additive genetic variance present in young and, particularly, old birds, but little evidence of any genetic variance for birds of intermediate ages. These results demonstrate that age can have a very important influence on the components of variation of characters in natural populations, and consequently that separate age classes cannot be assumed to be equivalent, either with respect to their evolutionary potential or response.

Organochlorines and mercury in waterfowl harvested in Canada.

Samples of breast muscle from 32 species of waterfowl collected from 123 sites across Canada were analyzed for chlorobenzenes (CBz), chlordane-related compounds (CHL), hexachlorocyclohexanes (HCH), DDT, mirex, dieldrin, PCBs and mercury. SigmaDDT, SigmaCBz and SigmaPCB were the compounds most frequently found above trace levels. SigmaHCH and SigmaMirex were detected the least often. Mercury was detected in all of the mergansers, over 50% of dabbling, bay and sea ducks, and in less than 2% of the geese analysed. The highest levels of contaminants were generally found in birds feeding at higher trophic levels such as sea ducks and mergansers. With the exception of a few samples of mergansers and long-tailed ducks from eastern Canada, which contained SigmaPCB concentrations of 1.0-2.4 mg kg(-1), SigmaPCB levels were less than 1 mg kg(-1) wet weight. Only one merganser from eastern Canada had a SigmaDDT concentration (2.6 mg kg(-1) ww) which was greater than 1 mg kg(-1) ww. The highest SigmaCHL (0.10 mg kg(-1) ww) was also found in mergansers from eastern Canada. Levels of total mercury in breast muscle were either low (< 1 mg kg(-1) ww) or below detection limits with the exception of a few samples of mergansers from eastern Canada which contained mercury concentrations of 1.0-1.5 mg kg(-1) ww. Health Canada determined that the organochlorine and mercury levels found in samples of breast muscle of ducks and geese analysed in this study did not pose a health hazard to human consumers and therefore these waterfowl were safe to eat.