Human–Wildlife Interactions 4(2):192–201, Fall 2010 Movements, habitat selection, associations, and survival of giant Canada goose broods in central Tennessee
1 ERIC M. DUNTON, Department of Biology, Tennessee Technological University, 1100 N. Dixie Avenue, Box 5063, Cookeville, TN 38505, USA [email protected] DANIEL L. COMBS, Department of Biology, Tennessee Technological University, 1100 N. Dixie Avenue, Box 5063, Cookeville, TN 38505, USA
Abstract: The brood-rearing period in giant Canada geese (Branta canadensis maxima) is one of the least-studied areas of goose ecology. We monitored 32 broods in Putnam County, Tennessee, from the time of hatching through fledging (i.e., when the goslings gained the ability to fly) and from fledging until broods left the brood-rearing areas during the spring and summer of 2003. We conducted a fixed-kernel, home-range analysis for each brood using the Animal Movement Extension in ArcView® 3.3 GIS (ESRI, Redlands, Calif.) software and calculated 95% and 50% utilization distributions (UD) for each brood. We classified 25 broods as sedentary (8 ha 95% UD), three as shifters (84 ha 95% UD), two as wanderers (110 ha 95%UD); two were unclassified because of low sample size. We measured 5 habitat variables (i.e., percentage of water, percentage of pasture, percentage of development, number of ponds, and distance to nearest unused pond) within a 14.5-ha buffer at nesting locations. We used linear regression, using multi-model selection, information theoretic analysis, to determine which, if any, habitat variables influenced home-range size at a landscape level. The null model was the best information-theoretic model, and the global model was not significant, indicating that landscape level habitat variables selected in this study cannot be used to predict home- range size in the Upper Cumberland region goose flock. We analyzed associations among broods, using a coefficient of association of at least 0.50, and determined association areas by overlaying individual home ranges. Overall gosling survival (Ŝ) during the brood-rearing period was 0.84 (95% CL = 0.78, 0.92), using a staggered-entry Kaplan-Meier survival curve. We believe that abundance of quality forage and pond habitat, high survivorship, and a lack of movement corridors (i.e., rivers, lakes, and reservoirs) were responsible for the relatively small home ranges of geese in the Upper Cumberland region. Associations formed during brood rearing may reduce predation risks and serve as a template for lifelong social bonds with family members and unrelated geese that are reared in the same locations.
Key words: brood associations, brood movements, Branta canadensis, Canada goose, home range, human–wildlife conflicts, survival, Tennessee
G C (Branta canadensis movements and habitat use (Eberhardt et al. maxima) were established across the United 1989). Previous approaches to study movements States and Canada in the 1960s and 1970s, and habitat use by broods have included color‑ primarily to provide hunting opportunities marking goslings (Geis 1956, Culbertson et al. where migratory populations had declined 1971), neck‑banding adults (Martin 1964, Zicus or never existed (Fritzell and Soulliere 2004, 1981, Mercer 1999), and placing radiotelemetry Griggs and Black 2004). The U.S. Fish and transmi ers on goslingsʹ parents (Lebeda and Wildlife Service estimates that 3.6 million giant Ra i 1983, Eberhardt et al. 1989, Didiuk and Canada geese exist in the United States (Haas Rusch 1998). Movement data of broods provide 2002), with the Mississippi Flyway supporting information about the distribution of problem the largest number (Nelson and Oe ing 1998). geese during critical periods (i.e., summer) and Increasing populations of giant Canada geese provide an estimate of productivity, which can have resulted in an increase in the number of be used to determine changes in flock density. human–goose conflicts. Understanding the Our objectives were to estimate home‑range ecology of giant Canada geese plays a crucial size and movement pa erns, test the influence role in managing urban–suburban problems. of landscape‑level habitat variables on home‑ Brood‑rearing is among the least understood range size, determine gosling survival rates, areas of goose ecology, particularly brood document the extent and circumstances
1Present address: Minnesota Department of Natural Resources, 35365 800th Avenue, Madelia, MN 56062, USA Canada goose broods • Dunton and Combs 193 surrounding formation of brood associations, broods occurred in areas not used by other and be er understand their function and effect broods, and the number and size of goslings on home‑range size and movements. were consistent among observations, providing strong evidence that we observed the same Study area broods despite the lack of individual marks. We conducted this study in the Highland Rim We observed broods daily or every other day province of Putnam County, Tennessee (Van throughout the brood‑rearing period. West 1998). Of the 105,198 ha in Putnam County, We established a driving route for brood 40% was farmland (mostly pastures), 40% was observations that began at the last known forestland, and 20% was urban environment location of broods. We recorded detailed (Van West 1998). National Wetland Inventory descriptions of brood locations, including the maps indicated that there are 2,292 palustrine distance from nearest major landmarks (e.g., open‑water habitats in Putnam County, and ponds, houses, or roads), time of observations, most were farm ponds (<1 ha). There were 3 and indications of disturbance (e.g., dogs, large water bodies in Putnam County: Boring mowing, cu ing, or other human activities). If Pond (14 ha), Cane Creek Lake (23 ha), and Old we did not locate broods at previously‑observed City Lake reservoir (37 ha). We did not include sites, we searched all nearby suitable habitats these large water bodies in this study because and ponds in increasing concentric circles either they were not representative of the common until we located broods or the search became nesting habitat types within the study area (i.e., too time‑consuming (i.e., we searched several 2 farms ponds <1 ha and pastures) and previous km ). We established an a priori sample size studies have documented disrupted nesting of 30 to 50 observations per brood to provide pa erns at one of these locations because of high accurate home‑range calculations (Seaman nesting densities (Mukherjee 2001, Christensen et al. 1999, Millspaugh and Marzluff 2001). 2002, White 2002). Although records were not If we did not locate a brood at least once in 7 available for verification, we suspected that the consecutive days, it was classified as missing, Upper Cumberland (UC) flock was established and we restricted searches to once a week at last in the late 1970s when birds were released on known locations and surrounding habitat. We farm ponds in the region, perhaps on the Boring considered broods absent a er a month of once‑ Pond (E. L. Warr, Tennessee Wildlife Resource a‑week searches failed (i.e., all goslings died) or Agency, personal communication). The best broods had moved sufficiently far enough away estimate of the size of the giant Canada goose that they could not be located. population in the UC region was 1,233 (White We observed broods once a week from the 2002). time of capture until they flew from their pre‑ fledging home ranges to determine how long Methods they utilized brood‑rearing areas once they We monitored 32 broods during spring and gained the ability to fly. If we were unable to summer 2003 from hatching through fledging locate broods during post‑fledging searches, (i.e., when the goslings gained the ability to we searched surrounding areas (i.e., all ponds fly) and from fledging until broods flew away and pastures within a few square kilometers of from rearing areas. We assumed that goslings last known locations) for 2 consecutive days. fledged at 70 days (Yocom and Harris 1965). We assumed broods not observed during these We obtained nesting locations, clutch sizes, searches had le brood‑rearing areas. and hatch dates from Carbaugh (2004), who conducted a concurrent nesting ecology Home-range analysis study. We monitored all pairs known to have We conducted home‑range analyses by successfully hatched ≥1 gosling within Putnam plo ing daily brood locations on digital‑ortho‑ County except for geese using the 3 large quarter quads (DOQQ) within ArcView 3.3 water bodies. At least 1 parent in 29 broods GIS so ware (ESRI, Redlands, Calif.). We used was marked with a white neck collar bearing a fixed‑kernel home‑range estimator for each a unique black, 4‑digit alphanumeric code, but brood using 50 and 95% utilization distributions 3 broods were completely unmarked. These (UD). We selected the Least Squares Cross 194 Human–Wildlife Interactions 4(2)
Validation (LSCV) as the smoothing parameter rearing period. To do so, we overlaid home (Seaman et al. 1999, Millspaugh and Marzluff ranges of all broods in that association, and 2001). digitized polygons around overlapping areas. We did not conduct fixed‑kernel home‑range Habitat analysis estimates for brood associations because brood We used linear regression to determine if sample sizes were inadequate (i.e., <30), thereby landscape level variables could predict home‑ violating a critical assumption (Millspaugh range size. We plo ed nesting locations on and Marzluff 2001). We calculated coefficients DOQQ maps, and generated circular buffers of of association (Cole 1949) for all brood 215‑m radii, which is equivalent to mean home‑ associations to determine how o en they were range size 14.5 ha, in ArcView and centered intact. We calculated coefficients of association at each nest. Within each buffer, we digitized by dividing the sum number of observations of area polygons to determine the percentage of broods when associated with other broods by 3 habitat types: water (e.g., ponds), pastures the sum total number of observations of those and lawns, and development (e.g., buildings same broods. Coefficients of associations >0.5 and roads). We calculated the number of ponds (i.e., 50% associations) are generally considered occurring within each buffer and the distance to be ecologically meaningful (Knight 1970, from each nest site to the nearest pond outside Millspaugh and Marzluff 2001). During the buffer not used by that brood. We employed individual observations, we considered broods a linear regression analysis that included away from their association if no other broods percentage of water, percentage of pasture, were in the immediate vicinity. However, we percentage of development, number of ponds, did not treat sightings impaired by nearby and distance to the nearest unused pond to landforms (e.g., hills or trees) as indications predict home‑range size. We used information of disassociation. We considered associations theoretic analysis for selecting models of all consisting of 3 or more broods intact if at least 2 possible combinations of independent variables broods were together at an observation. (Burnham and Anderson 1998). In addition, we generated both a null model that included no Survival analysis regression variables and a global model that We calculated gosling survival for the entire contained all variables (Long 1997). We selected brood‑rearing period using a staggered entry the model with the lowest biased‑corrected Kaplan‑Meier survival curve (Kaplan and Meier Akaike Information Criterion (AICc), and 1958). We discovered 5 broods post‑hatch that considered all models within 2 AICc points of had unknown nesting locations. We determined the best model as competing models (Burnham back‑dated hatch dates using gosling plumage and Anderson 1998). We excluded 6 broods in characteristics (Yocom and Harris 1965). We this analysis because 5 broods had unknown assigned the mid‑point between observation nesting locations, and we considered 1 brood dates as the mortality date for mobile broods an outlier because of an excessively large home and broods that we did not observed daily (n range, probably related to access to the Falling = 11). We conducted a sensitivity analysis to Water River that served as a movement corridor. determine the importance of these assumptions, We used Statistical Analysis System (SAS by perturbing unknown (estimated) hatch Institute, Cary, N.C.) for all statistical tests. dates ±3 days and se ing unknown mortality dates to the le and right endpoints of each Brood associations observation interval. We conducted a cluster‑ We defined brood associations as groups level bootstrap using 5,000 bootstrap samples of broods that were commonly near each to account for correlated survival within a other and exhibited synchrony in movements. brood. We used the 0.025 and 0.975 quantiles However, individual broods were usually of the bootstrap distribution to produce a 95% distinguishable within associations. We confidence interval for the Kaplan and Meier calculated brood association areas for broods (1958) estimate. that joined other broods during the brood‑ Canada goose broods • Dunton and Combs 195
Results Falling Water River (i.e., a movement corridor) Home-range analysis and had the largest home range in the study We recorded 1,340 brood observations (138.5 ha). The average age that broods le from April 21, 2003, to July 29, 2003, and brood‑rearing areas was 76.5 days, indicating drove >8,000 km along the observation route. that broods leave brood‑rearing areas soon Brood movement pa erns and home ranges a er they gain the ability to fly (Table 1). demonstrated 3 pa erns of movement (i.e., sedentary, shi ers, and wanderers), which Habitat analysis Hughes et al. (1994) described as: (1) sedentary Mean habitat within 14.5‑ha buffers around 26 broods that had 1 small, well‑defined area goose nests consisted of 58% pasture or lawns, of activity; (2) shi ers that commonly had 2 32% woods, 7% development, and 3% water. Table 1. Movement pa ern and home‑range size for the brood‑rearing period for 32 giant Canada goose broods in the Upper Cumberland region, Tennessee, 2003.
95% Utilization distribution 50% Utilization distribution
Number of SE Movement pattern broods 0 (ha) 0 (ha) SE
a Sedentary 25 7.6 1.5 1.6 0.3 b Shi er 3 84.0 17.6 15.0 3.9 c Wanderer 2 109.9 28.6 19.1 1.9 d Unclassified 2 7.6 0.7 1.7 0.3 a Sedentary broods had 1 small, well-defined area of activity. b Shifters commonly had 2 distinct areas of concentrated use occupied sequentially over the course of the brood- rearing season. c Wanderers ranged widely, having poorly-defined areas of use and no concentrated center of activity. d Two broods went missing during the brood-rearing season but appeared later.
Table 2. Best information theoretic habitat model Mean number of ponds was 2.3, and mean and the 3 competing models (i.e., within 2 AIC distance to the nearest pond not used was 429 points) for giant Canada geese in the Upper m. The global model to predict home‑range size Cumberland region, Tennessee. using all variables was not significant (P = 0.37, 2 Models R2 AICa R = 0.22). The best approximating model was Null 145.5257 the null model (AICc = 145.52), which contains Development 0.0568 146.3594 no regression variables. Only 3 models were Distance to pond 0.0524 146.4804 within 2 AICc points and considered competing (Table 2). There was a weak relationship Development + Distance to pond 0.1542 146.0945 between home‑range size and the amount of development and distance to the nearest pond a AIC corrected for small sample size. not used by the brood, but we determined no other measured variables to be important. distinct areas of concentrated use occupied sequentially over the course of the brood‑ Brood associations rearing season; and (3) wanderers that ranged Eighteen broods formed an association with widely, having poorly‑defined areas of use, and ≥1 other broods during the brood‑rearing no concentrated center of activity. We classified period, with a mean gosling age of 15 days (SE 25 broods as sedentary, three as shi ers, and = 3.6) at the time of group formation. In most two as wanderers. We did not classify 2 broods cases, brood associations formed immediately due to low sample size. Mean 95% utilization a er hatch and consisted of broods from nests distribution for sedentary broods was <10% the in close proximity to each other. Mean 95% UD size of the home range of shi er broods (Table for sedentary and broods that shi ed brood‑ 1). One of the wanderer broods had access to the rearing areas were similar, and coefficients of 196 Human–Wildlife Interactions 4(2)
Table 3. Movement pa ern, number of giant Canada goose brood associations, association home range size, and coefficient of association in the Upper Cumberland region, Tennessee during summer 2003.
a a Number of broods Coefficient of b Movement pa ern associated 95% UD (ha) 50% UD (ha) association Sedentary 15 8.1 1.6 0.88 (0.96) Shifters 3 8.6 2.6 0.79 (0.90) Wanderers 0 a UD = utilization distributions. We determined association home ranges by overlaying each brood in the as- sociation on each other and creating area polygons around overlapping areas. We did not conduct kernel home range estimations for brood associations because brood association sample sizes were inadequate (i.e., <30) and would have resulted in an inflated home-range estimate. b Coefficient of association = sum of observations of broods when associated ÷ sum of total observations of broods. Association values in parentheses represent coefficients after associations were formed.
between 33 and 41 days post‑hatch), indicating that all gosling mortality occurred within the first 5 weeks post‑hatch, with most mortality occurring in the first 2 weeks. The overall shape and ending survival estimate were not sensitive to assumed hatch or mortality dates.
Discussion Home‑range sizes of waterfowl broods are influenced by 3 primary factors. Foremost, sufficient food must be available to young waterfowl to meet energetic and nutritional demands of initial rapid physical growth (Sedinger 1986, MacInnes 1998, Mowbray et al. 2002) and growth and replacement of feathers during initial molts, which occur simultaneously (Sedinger 1986). Second, brood ! Figure 1. Kaplan-Meier survival distribution for movements o en are influenced by predation the brood-rearing period for 32 Canada goose risks because young waterfowl are highly broods in the Upper Cumberland region, Ten- vulnerable due to their small size and inability nessee, 2003. to fly (Ball et al. 1975, Talent et al. 1983, Rotella and Rath 1992). In addition, social interactions association were similar (Table 3). The 3 broods among broods, especially in geese, may that shi ed brood‑rearing areas and formed a contribute to movement pa erns. We believe brood association moved to a communal brood‑ Canada goose brood movements in the UC rearing area that was >1 km overland from their region are influenced by all of these factors and nest sites (Tables 2 and 3). the interaction effects among them. Optimal brood‑rearing habitat for Canada Survival analysis geese consists of gently sloping banks, nearby One hundred fi y‑six goslings hatched in the water reserves, few disturbances, and abundant study area, and 132 goslings fledged successfully. plant food in the form of short grasses, semi‑ Overall survival (Ŝ) during the brood‑rearing aquatic plants, or emergent vegetation (Hanson period was 0.84 (95% CL = 0.78, 0.92; Figure 1). and Eberhardt 1971, Bellrose 1980, Sedinger Eighteen of the 32 broods successfully fledged and Raveling 1986, Bruggink et al. 1994). all goslings, and only 5 broods lost >50% of their Foraging habitat in the UC appears to be goslings. We did not detect changes in gosling evenly distributed in sufficient quantities to numbers a er approximately 5 weeks (i.e., meet growth demands on goslings, perhaps Canada goose broods • Dunton and Combs 197 explaining why proportion of land in pastures size and shi ing or wandering movement was not linked to home‑range size. pa erns for some broods in this study, but Habitat conditions influence movement only a er goslings reached the critical age that pa erns and home‑range size in waterfowl and reduced predation risks (i.e., 4 to 5 weeks). many other animals (Eberhardt et al. 1989, Dzus Parents may form brood associations with and Clark 1997, Didiuk and Rusch 1998, Yerkes siblings or with adults with which they were 2000). Movements and or increased home ranges previously associated. Canada goose broods are sometimes caused by a deficiency of specific o en associate with other broods, sometimes habitat requirements within a concentrated or moving from their natal area to communal localized area (Mauser et al. 1994, Mizutani brood‑rearing areas (Geis 1956, Zicus 1981, and Jewell 1998, and Yerkes 2000). Regression Eberhardt et al. 1989, Didiuk and Rusch analysis conducted in this study indicates that 1998). Although li le is known about social portions of Putnam County used by nesting interactions during brood‑rearing (Mulder et al. geese are relatively uniform in habitat condi‑ 1995), various theories have been proposed for tions. Sedentary broods used habitats similar to why geese form brood associations. Most o en those of geese that shi ed brood‑rearing areas. cited explanations are dominance relationships Limited brood mobility observed in this study and competition for food among various‑sized was undoubtedly influenced by the landscape groups, reduction in predation risks, and of the study area. Sca ered farm ponds in inadvertent mixing of young among broods a rural–suburban se ing provide excellent (Gosser and Conover 1999). Brood associations brood‑rearing habitat, but on a localized may simply reflect brood‑site fidelity by parents basis. Shortage of water corridors necessitated (i.e., use of the same location by several broods; overland travel, and parents appeared reluctant Zicus 1981, Didiuk and Rusch 1998, Lindberg to move their broods, especially during the first and Sedinger 1998). Although advantages of few weeks post‑hatch. Brood movements and belonging to brood associations are not fully fidelity will vary in different landscapes, as understood, short‑term benefits to young in shown by greater movements along shorelines close family associations seem clear: they are of reservoirs (Eberhardt et al. 1989, Mercer a acked less o en, feed more, and have access 1999). The brood with the largest home range to food and space in relation to the dominance in this study (138.5 ha) used a water corridor, status of their parents (Raveling et al. 2000). supporting this conclusion. Many urban goose problems are associated Because habitat was relatively uniform, we with large congregations of geese throughout believe differences in movement pa erns and the summer, and the propensity for broods home‑range size among broods in this study to associate contributes to the problem. Most were influenced by other factors more than broods in the UC region formed associations foraging habitat. If Canada geese survive their soon a er hatching, but they generally first year, their annual survival rates become dispersed from brood‑rearing areas within higher, a key reason for exponential growth of a week of fledging, providing evidence that urban flocks (Smith et al. 1999). Survival rates management activities should be utilized prior of other Canada goose flocks varied from 5 to to nest initiation. 95% during the brood‑rearing period (Martin A key component of any management 1964, Combs et al. 1984, Baker 1989, Huskey strategy is monitoring and evaluation. Due et al. 1998). High survival rates (i.e., 84%) in to the reduced mobility during the brood‑ this study indicate that there is low predation rearing period, Canada goose broods are easily risk and that survival during the brood‑rearing observed, providing a mechanism both to period is not limiting population growth in the easily index annual productivity and determine UC region. changes in flock density. In Georgia, Powell et al. Canada geese are highly social and have a (2004) used a postcard survey of golf courses to well‑developed social system (Raveling 1969, monitor urban subpopulations of Canada geese 1970; Combs 1989; Christensen et al. 2002). and found the technique to be a cost‑effective Parental desire to form brood associations tool to provide information on a segment of the probably contributed to increased home‑range population that is hard to quantify with other 198 Human–Wildlife Interactions 4(2) techniques. A well‑designed brood survey are considered undesirable (e.g., golf courses, based on a thorough understanding of Canada parks, and manicured lawns). Most landowners goose brood‑rearing ecology (e.g., movement in the UC region have a rural background and pa erns, social biology, and site fidelity) can have expressed li le animosity toward geese provide biologists with critical information unless they accumulate in large numbers. Many in a timely and relevant manner (Powell et al. landowners are protective of geese, especially 2004). Because most geese exhibit a high degree broods that are reared on their property. of site fidelity and most urban goose conflicts However, many farms in the region are now are highly localized, a brood route survey of being sold and subdivided, and homeowners in brood‑rearing areas would provide an index subdivisions generally are less tolerant of geese. of changes in flock density and evidence of Fidelity of geese to specific ponds is likely to effectiveness of management activities. cause future conflicts in a changing landscape. Urban goose problems are o en complex Such changes should be considered when and involve an integrated management developing management strategies, especially approach that provides short‑term and long‑ if they involve releasing or translocating geese. term solutions to managing goose populations at or below target levels (Smith et al. 1999). Acknowledgments Urban populations are o en difficult to survey We thank J. Fieberg, wildlife biometrician, and monitor because they are widespread and Minnesota Department of Natural Resources, sca ered (Powell et al. 2004). Although human for providing assistance with the survival populations in the UC region are not directly analysis. S. Haysle e reviewed early dra s of comparable with large metropolitan areas, the manuscript and provided input on analysis. habitat conditions (e.g., numerous ponds and We also thank the private landowners in abundant grasslands) are reflective of giant Putnam County who allowed us to access their Canada goose habitat in many suburban areas land for this study. Funding for this project was that experience goose problems. 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Raveling, D. G. 1970. Dominance relationships ERIC M. DUNTON is a wildlife research and agnostic behavior of Canada geese in win- biologist with Minnesota Department of Natural ter. Behaviour 37:291–319. Resources. His current research Raveling, D. G., J. S. Sedinger, and D. S. Johnson. involves ecology 2000. Reproductive success and survival in re- and manage- ment of wild tur- lation to experience during the first two years in keys and turkey Canada geese. Condor 102:941–945. hunt manage- ment. He re- Rotella, J. J., and J. T. Rath. 1992. Mallard brood ceived his B.S. movements and wetland selection in south- degree from western Manitoba. Journal of Wildlife Manage- Michigan State University and ment 56:508–515. his M.S. degree Seaman, D. E., J. J. Millspaugh, B. J. Kernohan, G. from Tennessee Technological C. Brundige, K. J. Raedeke, and R. A. Gitzen. University and 1999. Effects of sample size on kernel home is a certified as- range estimates. Journal of Wildlife Manage- sociate wildlife biologist with The Wildlife Society. ment 63:739–747. Sedinger, J. S. 1986. Growth and development of DANIEL L. COMBS is a professor and the Canada goose goslings. Condor 88:169–180. chair of the Department of Biology at Tennessee Sedinger, J. S., and D. G. Raveling. 1986. Timing Technological University. of nesting by Canada geese in relation to the His primary research interest is ecology and phenology and availability of their food. Journal management of water- of Animal Ecology 55:1083–1102. fowl, especially Canada geese. He received his Smith, A. E., S. R. Craven, and P. D. Curtis. 1999. B.S. and M.S. degrees Managing Canada geese in urban environ- from Auburn University ments. Jack Berryman Institute and Cornell and his Ph.D. degree from the University of University Cooperative Extension Publication Missouri. He is a certi- 16, Ithaca, New York, USA. fied wildlife biologist and a past president of the Talent, L. G., R. L. Jarvis, and G. L. Krapu. 1983. Tennessee Chapter of Survival of mallard broods in south-central The Wildlife Society. North Dakota. Condor 85:74–78. Van West, C. 1998. The Tennessee encyclope- dia of history and culture. Rutledge Hill Press, Nashville, Tennessee, USA. White, H. B. 2002. Movement patterns and sub- flock associations of giant Canada geese in the Upper Cumberland region, Tennessee. Thesis, Tennessee Technological University, Cooke- ville, Tennessee, USA. Yerkes, T. 2000. Influence of female age and body mass on brood and duckling survival, number of surviving ducklings, and brood movements in redheads. Condor 102:926–929. Yocom, C. F., and S. W. Harris. 1965. Plumage descriptions and age data for Canada goose goslings. Journal of Wildlife Management 29:874–877. Zicus, M. C. 1981. Canada goose brood behavior and survival estimates at Crex Meadows, Wis- consin. Wilson Bulletin 93:207–217.