R. Michael Anthony

Feedback dynamics of grazing lawns: coupling vegetation change with animal growth

We studied the effects of grazing by Black Brant (Branta bernicla nigricans) geese (hereafter Brant) on plant community zonation and gosling growth between 1987 and 2000 at a nesting colony in southwestern Alaska. The preferred forage of Brant, Carex subspathacea, is only found as a grazing lawn. An alternate forage species, C. ramenskii, exists primarily as meadow but also forms grazing lawns when heavily grazed. We mowed plots of ungrazed C. ramenskii meadows to create swards that Brant could select and maintain as grazing lawns. Fecal counts were higher on mowed plots than on control plots in the year after plots were mowed. Both nutritional quality and aboveground biomass of C. ramenskii in mowed plots were similar to that of C. subspathacea grazing lawns. The areal extent of grazing lawns depends in part on the population size of Brant. High Brant populations can increase the areal extent of grazing lawns, which favors the growth of goslings. Grazing lawns increased from 3% to 8% of surface area as the areal extent of C. ramenskii meadows declined between 1991 and 1999. Gosling mass was lower early in this time period due to density dependent effects. As the goose population stabilized, and area of grazing lawns increased, gosling mass increased between 1993 and 1999. Because larger goslings have increased survival, higher probability of breeding, and higher fecundity, herbivore-mediated changes in the distribution grazing lawn extent may result in a numerical increase of the population within the next two decades.

FOODS OF ARCTIC FOXES (ALOPEX LAGOPUS) DURING WINTER AND SPRING IN WESTERN ALASKA

Abstract During 1986–1991, carcasses of 619 arctic foxes (Alopex lagopus) collected from local trappers and at biological field camps on the Yukon-Kuskokwim Delta in western Alaska from November through May were analyzed to determine gastrointestinal contents, age, sex, and body condition. Prey in declining order of importance were small mammals (95% tundra voles, Microtus oeconomus), birds, marine mammals, and fishes. Foxes with small mammal remains in their stomachs were captured farther from the Bering Sea coast (X̄ = 5.2 km) than those without small-mammal remains (2.8 km); foxes consuming remains of marine mammals were closer to the coast (1.9 km) than others (4.9 km). Although eggshells had a poor likelihood of occurrence in stomachs, they were found in all months and years. In 1986 and 1987, foxes consumed fewer small mammals than in other years. Mean ages of foxes captured in 1986 (3.7 years) and 1987 (3.2) were greater than in all other years (1.5). Capture of adults was more common as winter progressed. Indexes of subcutaneous fat decreased annually in April–May and were highest in 1991, when occurrence of carrion of marine mammals was highest.

OPTIMIZING NEST SURVIVAL AND FEMALE SURVIVAL: CONSEQUENCES OF NEST SITE SELECTION FOR CANADA GEESE

Abstract We examined the relationship between attributes of nest sites used by Canada Geese (Branta canadensis) in the Copper River Delta, Alaska, and patterns in nest and female survival. We aimed to determine whether nest site attributes related to nest and female survival differed and whether nest site attributes related to nest survival changed within and among years. Nest site attributes that we examined included vegetation at and surrounding the nest, as well as associations with other nesting birds. Optimal nest site characteristics were different depending on whether nest survival or female survival was examined. Prior to 25 May, the odds of daily survival for nests in tall shrubs and on islands were 2.92 and 2.26 times greater, respectively, than for nests in short shrub sites. Bald Eagles (Halieaeetus leucocephalus) are the major predator during the early breeding season and their behavior was likely important in determining this pattern. After 25 May, when eagle predation is limited due to the availability of alternative prey, no differences in nest survival among the nest site types were found. In addition, nest survival was positively related to the density of other Canada Goose nests near the nest site. Although the number of detected mortalities for females was relatively low, a clear pattern was found, with mortality three times more likely at nest sites dominated by high shrub density within 50 m than at open sites dominated by low shrub density. The negative relationship of nest concealment and adult survival is consistent with that found in other studies of ground-nesting birds. Physical barriers that limited access to nest sites by predators and sites that allowed for early detection of predators were important characteristics of nest site quality for Canada Geese and nest site quality shifted within seasons, likely as a result of shifting predator-prey interactions.

Nest survival in dusky Canada geese ( Branta canadensis occidentalis ): Use of discrete-time models

Abstract The Dusky Canada Goose (Branta canadensis occidentalis) population that breeds in the Copper River Delta, Alaska, has declined substantially since the late 1970s. Persistent low numbers have been attributed to low productivity in recent years. We examined patterns in survival rates of 1,852 nests to better understand ecological processes that influenced productivity during 1997-2000. We compared 10 nonparametric models of daily survival rate of nests (DSR) that included variation among years, calendar dates, nest initiation dates, and nest ages with equivalent models based on parametric functions. The unequivocal best model included patterns of DSR that varied among discrete periods of years, calendar dates, and nest ages. Generally, DSR was low early in the nesting season and higher midseason. Across years, patterns in DSR were most variable early and late in the nesting season. Daily survival rates of nests declined between the first and second week after initiation, increased until the fourth week, and then declined during the last week before hatch. Nest survival probability estimates ranged from 0.07 to 0.71 across years and nest initiation dates. Mean rates of nest survival ranged between 0.21 and 0.31 each year. We suggest (1) considering models that do not limit estimates of daily nest survival to parametric forms; (2) placing greater emphasis on sample size when nests are rare, to obtain accurate estimates of nest survival; and (3) developing new techniques to estimate the number of nests initiated. Supervivencia de los Nidos en Branta canadensis occidentalis: Uso de Modelos de Tiempo Discreto

A quantitative approach to identifying predators from nest remains

Abstract Nesting success of Dusky Canada Geese (Branta canadensis occidentalis) has declined greatly since a major earthquake affected southern Alaska in 1964. To identify nest predators, we collected predation data at goose nests and photographs of predators at natural nests containing artificial eggs in 1997–2000. To document feeding behavior by nest predators, we compiled the evidence from destroyed nests with known predators on our study site and from previous studies. We constructed a profile for each predator group and compared the evidence from 895 nests with unknown predators to our predator profiles using mixture-model analysis. This analysis indicated that 72% of destroyed nests were depredated by Bald Eagles and 13% by brown bears, and also yielded the probability that each nest was correctly assigned to a predator group based on model fit. Model testing using simulations indicated that the proportion estimated for eagle predation was unbiased and the proportion for bear predation was slightly overestimated. This approach may have application whenever there are adequate data on nests destroyed by known predators and predators exhibit different feeding behavior at nests.

Techniques for identifying predators of goose nests

Abstract We used cameras and artificial eggs to identify nest predators of dusky Canada goose Branta canadensis occidentalis nests during 1997–2000. Cameras were set up at 195 occupied goose nests and 60 artificial nests. We placed wooden eggs and domestic goose eggs that were emptied and then filled with wax or foam in an additional 263 natural goose nests to identify predators from marks in the artificial eggs. All techniques had limitations, but each correctly identified predators and estimated their relative importance. Nests with cameras had higher rates of abandonment than natural nests, especially during laying. Abandonment rates were reduced by deploying artificial eggs late in laying and reducing time at nests. Predation rates for nests with cameras were slightly lower than for nests without cameras. Wax-filled artificial eggs caused mortality of embryos in natural nests, but were better for identifying predator marks at artificial nests. Use of foam-filled artificial eggs in natural nests was the most cost effective means of monitoring nest predation.

Survival of Dusky Canada Goose Goslings in Relation to Weather and Annual Nest Success

Abstract The dusky Canada goose (Branta canadensis occidentalis) population has been in long-term decline, likely due to reduced breeding productivity, but gosling survival of this population had not been examined. We studied gosling survival in broods of radiomarked adult females on the western Copper River Delta, Alaska, USA, during 1997–1999 and 2001–2003. Survival estimates for dusky Canada goose goslings to 45 days (x̄ = 0.32) were below estimates from most previous studies of geese. Daily survival of goslings increased with age and decreased with date of hatch. Precipitation during the first 3 days post-hatch was negatively related to gosling survival and this effect increased with date. Annual estimates of gosling survival were positively correlated with annual estimates of nest success, suggesting overlap in factors affecting nest and gosling survival. Nest success probably also directly affected gosling survival, because survival decreased with hatch date and more broods hatched from renests during years with low nest success. Gosling survival appears to play an important role in limiting current productivity of this population. Management directed at increasing nest success would likely also improve gosling survival. We recommend additional research directed at examining sources of gosling mortality and the link between nest success and gosling survival.

Renesting by Dusky Canada Geese on the Copper River Delta, Alaska

Abstract The population of dusky Canada geese (Branta canadensis occidentalis; hereafter duskies) breeding on the Copper River Delta (CRD), Alaska, USA, has been in long-term decline, largely as a result of reduced productivity. Estimates of renesting rates by duskies may be useful for adjusting estimates of the size of the breeding population derived from aerial surveys and for understanding population dynamics. We used a marked population of dusky females to obtain estimates of renesting propensity and renesting interval on the CRD, 1999–2000. Continuation nests, replacement nests initiated without a break in the laying sequence, resulted only after first nests were destroyed in the laying stage with ≤4 eggs laid. Renesting propensity declined with nest age from 72% in mid-laying to 30% in early incubation. Between first nests and renests, mean interval was 11.9 ± 0.6 days, mean distance was 74.5 m (range 0–214 m), and clutch size declined 0.9 ± 0.4 eggs. We incorporated our renesting estimates and available estimates of other nesting parameters into an individual-based model to predict the proportion of first nests, continuation nests, and renests, and to examine female success on the CRD, 1997–2000. Our model predicted that 19–36% of nests each year were continuation nests and renests. Also, through 15 May (the approx. date of breeding ground surveys), 1.1–1.3 nests were initiated per female. Thus, the number of nests per female would have a significant, though relatively consistent, effect on adjusting the relation between numbers of nests found on ground surveys versus numbers of birds seen during aerial surveys. We also suggest a method that managers could use to predict nests per female using nest success of early nests. Our model predicted that relative to observed estimates of nest success, female success was 32–100% greater, due to replacement nests. Thus, although nest success remains low, production for duskies was higher than previously thought. For dusky Canada geese, managers need to consider both continuation nests and renests in designing surveys and in calculating adjustment factors for the expansion of aerial survey data using nest densities.

Snow cover and snow goose Anser caerulescens caerulescens distribution during spring migration

Arctic geese often use spring migration stopover areas when feeding habitats are partially snow covered. Melting of snow during the stopover period causes spatial and temporal variability in distribution and abundance of feeding habitat. We recorded changes in snow cover and lesser snow goose Anser caerulescens caerulescens distribution on a spring migration stopover area in south-central Alaska during aerial surveys in 1993–1994. Our objectives were to determine whether geese selected among areas with different amounts of snow cover and to assess how temporal changes in snow cover affected goose distribution. We also measured temporal changes in chemical composition of forage species after snow melt. We divided an Arc/Info coverage of the approximately 210 km2 coastal stopover area into 2-km2 cells, and measured snow cover and snow goose use of cells. Cells that had 10–49.9% snow cover were selected by snow geese, whereas cells that lacked snow cover were avoided. In both years, snow cover diminished along the coast between mid-April and early May. Flock distribution changed as snow geese abandoned snow-free areas in favour of cells where snow patches were interspersed with bare ground. Snow-free areas may have been less attractive to geese because available forage had been quickly exploited as bare ground was exposed, and because soils became drier making extraction of underground forage more difficult. Fiber content of two forage species increased whereas non-structural carbohydrate concentrations of forage plants appeared to diminish after snow melt, but changes in nutrient concentrations likely occurred too slowly to account for abandonment of snow-free areas by snow geese.

Predation and reduced grazing interact to reduce recruitment and population growth in Black Brant

ABSTRACT Dynamics of avian populations may be governed by a complex interaction between immediate effects of predation and longer-term trophic feedbacks between individuals and their foods. We used a long-term study of uniquely marked Black Brant geese (Branta bernicla nigricans) to estimate recruitment into the breeding population. We related recruitment to nest success, which directly affects recruitment 2 to 3 years later. We also assessed similarly time-lagged relationships between recruitment and number of nests in the colony, a measure of local density, and pre-fledging and first-year survival, the latter of which is strongly influenced by growth conditions and food availability for young of the year. We assessed relationships between number of recruits and explanatory variables in 2 sets of models, which either included or excluded a linear trend across years in numbers of recruits. The best-performing models in each model set explained a substantial proportion of the variation in numbers of recruits 2 to 3 years later (85 and 78% in the 2 model sets, respectively). First-year survival was an important predictor of number of recruits in both sets of models, whereas nest success was important only in models lacking a trend across years. Number of nests in the colony had a relatively weak negative association with number of recruits. First-year survival reflects a lagged response to earlier nest-predation events. Reduced grazing during predation events can result in loss of grazing lawns and thus reduced growth rates of future cohorts of goslings, which are regulated by food abundance and density of broods. Our finding that recruitment is insufficient to maintain the breeding population results from both direct effect of predation on nests, primarily by Arctic foxes (Alopex lagopus), and longer-term indirect effects of such predation on first-year survival of Black Brant.