Reproductive success and brood movements of giant Canada geese in Eastern South Dakota.
The South Dakota Department of Game, Fish, and Parks (SDGFP) initiated a program to reintroduce giant Canada geese (Branta canadensis maxima) into the state in 1962 (Mammenga, 2000). During these efforts, several research projects were conducted to examine reproductive success of these geese (Hilley, 1976; Clausing, 1979). In recent years, South Dakota's resident Canada goose population has increased dramatically. The 2005 May breeding population in the eastern half of South Dakota was estimated at 108,000 (U. S. Fish and Wildlife Service, 2006). As in many other areas of the United States, there have been problems associated with high populations of Canada geese. In order to properly manage giant Canada goose populations as they increase, it is important to determine their reproductive success. During an ongoing study of giant Canada goose movements (Anderson, 2006), we had the opportunity to determine the reproductive success of radio-collared females. We wanted to determine reproductive parameters such as clutch sizes, nest initiation dates, hatching dates, and nesting success since the reintroduction period.
In South Dakota, a problem caused by giant Canada geese is crop damage, especially in soybean fields (Schaible et al., 2005). In 1996, SDGFP initiated a program to reduce crop damage by giant Canada geese. This $200,000/yr program was created to provide landowners that file a complaint free access to abatement techniques (Mammenga, 2000). The program has been in use for over a decade, but landowner complaints continue to occur. Since most crop damage is caused by molting or locally raised geese (Schaible et al., 2005), biologists were interested in movements of adult females and their broods from nesting sites to brood-rearing sites. Of primary interest was determining how far adults moved their broods to reach summer molting wetlands because concentrations of young and molting geese lead to increased crop damage (Schaible et al., 2005). The objectives of this study were (1) to define nesting parameters of marked female giant Canada geese in eastern South Dakota including nesting phenology, nest success, hatching success, egg success, clutch size, and number of goslings leaving successful nests, and (2) to document and measure movements of adult female giant Canada geese and their broods from nesting sites to wetlands where the brood was raised and adults molted.
STUDY AREA AND METHODS
Northeastern South Dakota lies in the Coteau des Prairies (Coteau) physiographic region (Gab, 1979). The Coteau is a glaciated area within the Prairie Pothole Region with many perennial and intermittent lakes and wetlands that are used extensively by breeding and staging waterfowl (Johnson and Higgins, 1997; Hogan and Fouberg, 1998). The numerous wetlands provide prime nesting conditions for giant Canada geese. Native vegetation of the area is tall grass prairie, which lies along the eastern edge of South Dakota, and gradually gives way to the northern mixed grass prairie to the west. Because of the study area's agricultural productivity and rather level topography, the greatest portion of tall grass prairie has now been replaced by agricultural crops (Hogan and Fouberg, 1998). Major crops of this area are corn (Zea mays), soybeans (Glycine max), wheat (Triticum aestivum), barley (Hordeum vulgare) alfalfa (Medicago sativa), and other hay.
We captured adult giant Canada geese in late Jun. and early Jul, at 25 sites in Brookings, Clark, Codington, Day, Hamlin, Kingsbury, and Lake Counties during the summers of 2000-2003. During this period, adults were molting and flightless while goslings were incapable of flight. Geese were captured by driving them into a corral-type trap (Cooch, 1953) that was erected in a bay with a gradual shoreline. Collars with transmitters (total wt. 60 g) were attached to 5-10 adult breeding females (as evidenced by a brood patch) at each capture site. VHF transmitters were manufactured by Advanced Telemetry Systems, Inc. (Isanti, MN), and were attached to black neck collars made by Paul Mammenga (SDGFP). VHF transmitters transmitted continuously (pulse rate of 50 ppm and pulse width of 20 ms) at frequencies within the 150 and 151 MHz range and had a life expectancy of 12 mo. We collected reproductive data on geese that had functioning transmitters at the beginning of each spring nesting season. We used a 4-element null-peak antenna system mounted on top of a pickup and a handheld yagi antenna to locate marked geese. We monitored the reproductive status of marked females 1-2 times per week from their arrival in Mar. until 1 Jul. Geese were classified as nonbreeders (not observed to have initiated a nest), unsuccessful nesters, and successful nesters. It is possible, but unlikely, that some geese classified as nonbreeders attempted nest initiation without being detected. Unsuccessful nesters were geese that attempted nesting but failed to hatch at least one gosling, while successful nesters hatched at least one gosling.
We searched for nests weekly and when nest sites were located, we determined the status of the nest. We recorded the Universal Transverse Mercator (UTM) coordinates, nest type (island, vegetation mound, or shoreline), date, time, nest condition, and incubation stage (by floating eggs) for all located nests (Westerskov, 1950). When a marked female was located away from her nest, we recorded the UTM coordinates, presence/absence of a pair, territorial displays, and brood count (if a successful nester). The island nest type included natural islands and flooded stockdam berms, while nest types classified as vegetation mounds included muskrat (Ondatra zibethicus) huts, cattail (Typha spp.) mounds, and other vegetation piles. After inspecting the nests, we covered eggs with nest material and down to help keep eggs warm and to reduce the likelihood of human-induced depredation.
The nest initiation date was defined as the date the first egg was laid in a nest. The period from when the last egg was laid until the first gosling left the egg was defined as the incubation period. The nesting season was the period from when the first egg was laid until the last egg hatched. We determined the date of nest initiation by backdating from day of hatch the mean incubation period of 28 d and a laying interval of 1.5 d per egg (Kossack, 1950; Brakhage, 1965; Cooper, 1978). If no eggs from a nest hatched, we estimated the date of nest initiation by backdating from the first day of nest visit (Westerskov, 1950) and a laying interval of 1.5 d per egg.
We monitored nests of marked females until the nest was lost, abandoned, or eggs had hatched. Nest fates were classified as successful, predator destroyed, flooded, and deserted. A successful nest was defined as having at least one egg hatch. Successful nests either contained goslings or had egg shells and membrane characteristics of hatched eggs (Cooper, 1973). We assumed a nest had been destroyed by predators when the entire clutch was missing or the eggs were broken and scattered. Flooded nests were submerged or had signs of recently being flooded. Deserted nests were those that contained breast down and cold eggs (Cooper, 1973). We re-visited deserted nests to confirm that they were abandoned.
We estimated apparent and Mayfield (Mayfield, 1961, 1975) nest success for all monitored nests. Apparent nest success was calculated as the proportion of nests that hatched at least one egg (Klett and Johnson, 1982). Apparent nest success has been shown to overestimate true success due to potential biases and data loss (Mayfield, 1961, 1975; Miller and Johnson, 1978; Klett and Johnson, 1982). Since all geese in this study were radio-marked and were closely monitored as soon as they returned to South Dakota in spring, the assumption that successful and unsuccessful nests were discovered at an equal probability was probably met. We calculated Mayfield nesting success by raising the daily survival rate to a power equal to the incubation period. Chi-square tests were used to determine if there was a difference in nest success by nest type.
Egg success was defined as the percentage of eggs that hatched from all nests. Hatching success was the percentage of eggs hatched from only successful nests. We estimated overall egg and hatching success following Cooper (1978). The effects of nest type and year on egg and hatching success were analyzed using Analysis of Variance (SAS Institute, 2004), following tests to ensure normality.
The largest number of eggs known to have been laid in the nest was defined as the total clutch size (TCS). We measured TCS for both successful and unsuccessful nests, but not for dump nests (which contained large numbers of unattended eggs) or nests deserted prior to completion. Goslings leaving the nest (GLN) was defined as the number of goslings leaving the nest with the parents. The effects of nest type and year on TCS and GLN were analyzed using Analysis of Variance (SAS Institute, 2004), following tests to ensure normality. Means are given [+ or -] standard error, and an alpha = 0.05 was used for all tests. The relation between clutch size and day of nest initiation was analyzed using Pearson's r statistic (SAS Institute, 2004).
We monitored marked females weekly to document their movements from the beginning of the nesting period through the molting period. Using ArcView[R] GIS 3.2 software, we measured the distance from all successful nest sites to the edge of the wetland where the marked goose was captured the previous year to determine how far they were moving. We monitored broods weekly in order to document their movements to summer molting sites and their movements between wetlands. Due to the inaccessibility of many brood wetlands, we could not estimate gosling survival. For geese that nested on wetlands other than where they were captured, we recorded the date they returned to their capture wetland.
We monitored the reproductive stares of 88 females during 2001-2004. Nest initiation dates ranged from 25 Mar. in 2004 to 1 May in 2002. Most (43/61, 71%) nests were initiated in the first two weeks of April. The nesting season lasted 61, 53, and 60 d during springs 2001-2003, respectively. The earliest hatch date was 2 May 2004 and the latest was 2 Jun. 2001. Most (28/ 42, 67%) of the hatch dates were from May 10-20. We observed unmarked geese with hatchlings up to a week earlier and up to 10 d later than geese included in this study.
Of the 88 females monitored, half of the geese had successful nests, 20.5% were unsuccessful and 29.5% of the geese did not attempt to nest (Table 1). We found no evidence of renesting by the 18 marked geese that had failed initial nesting attempts. Most nests were on islands (36/62, 58%) followed by vegetation mounds (17/62, 27%) and shorelines (9/62. 14%). Mayfield and apparent estimates for mean nest success across years was 62.7% and 71.0%, respectively, (Table 2). There was no difference ([chi square] = 3.20, 2 df, P = 0.202) in nest success by nest type across years. Predation was the primary cause of nest failure (10/62, 16%) across years. As water levels retreated during spring 2003 and 2004 over much of the study area, water became shallow enough to allow predators to access many islands. We documented complete predation of all nests on three islands with marked geese nesting on them. The islands had approximately 30 other goose nests, which were all destroyed by predators within a 7-day period. Flooding destroyed seven nests during the springs of 2001 and 2002.
Overall mean egg success was 62.6 % (range 47-71%) with 223 of 356 eggs hatching. There was no difference in egg success by year ([F.sub.3,61] = 3.55, P = 0.49) or nest type ([F.sub.3,43] = 3.11, P = 0.77). Destruction by predators (63 eggs, 17%), flooding (41 eggs, 11%), and infertility (14 eggs, 4%) were the largest contributors to egg failure. The mean hatching success was 88.8% (223/251 eggs) from successful nests. There was no difference in hatching success by year ([F.sub.3,43] = 3.20, P = 0.25) or nest type ([F.sub.3,43] = 2.18, P = 0.48). Mean TCS across years was 5.73 [+ or -] 0.17 (n = 60), and ranged from two to nine eggs. There were no differences in TCS among nest types ([F.sub.2,59] = 1.02, P = 0.96,) or among years ([F.sub.3,59] = 2.05, P = 0.41,). There was no correlation between TCS and day of nest initiation for combined years (n = 60, [chi square] = 0.04, P = 0.147). We recorded one dump nest (11 eggs) for a marked goose. Mean number of GLN across years was 5.02 [+ or -] 0.25 (n = 44), and ranged from one to eight goslings. There was no difference among years in GLN ([F.sub.3,59] = 3.61, P = 0.261) or among nest types in GLN ([F.sub.3,59] = 1.44, P = 0.88).
Fifty-five VI-IF marked geese maintained functional transmitters up to the molting period. Twenty-seven (49.1%) marked females nested on or around the shoreline of their capture/ molt wetland. The remaining 28 (50.9%) nested on other wetlands and the mean distance from capture/molt wetlands to nest site wetlands was 1.5 km [+ or -] 0.18 (range 0.1-4.1 km).
Eighteen of 27 marked females that nested on their capture/molt wetland were successful. Nineteen of 28 marked geese that nested on wetlands other than their capture/ molt wetland were successful and all but 2 returned to their capture/molt wetlands from 6 May-14 Jun. Seven of these 19 geese returned to their capture/molt wetlands with their broods less than three days after hatching. Ten other geese returned to their capture/molt wetlands within one to five weeks after hatching. One goose remained at a seasonal wetland 2.8 km from its capture wetland raising a gang brood. Another goose moved with its brood 6.9 km overland through at least six seasonal and semipermanent wetlands. The mean number of wetlands used by the 19 successful geese with broods prior to reaching their molt wetland was 2.5 [+ or ] 0.23 (range 2-6). Generally, marked geese would move directly overland from their nesting wetland site to the molting wetland site. All marked geese that nested on their capture/molt wetlands remained there to molt and rear their broods. Some wetlands were mainly used for molting rather than nesting such as Goodfellow's wetland in Brookings County. Geese were captured from Goodfellow's wetland during 2001 and 2003, but no marked geese nested on this wetland. Geese that had a successful nest returned to Goodfellow's wetland to molt and rear their broods.
The nesting season length in eastern South Dakota was shorter than most other giant Canada goose nesting durations (Table 3), but we observed unmarked females initiating nests earlier and hatching goslings later than marked geese, indicating the true season was longer. Canada geese generally have a single nest, but renesting may occur under certain circumstances (Geis, 1956; Atwater, 1959; Brakhage, 1965; Lengkeek, 1973; Sayler, 1977; Cooper, 1978). We found no evidence of renesting attempts by marked females during this study, suggesting that renesting was of little importance in eastern South Dakota. Two marked geese that were on the study area prior to 1 April did not initiate their nests until late spring (26 April and 1 May). Cooper (1978) stated that renests can easily be identified by the lateness of their establishment, and some researchers (Nigus and Dinsmore, 1980) have used this criterion to identify renests. However, we caution researchers to use other criteria than lateness to classify a nest as a renest, since it may actually be a late original nest.
Although we did not know the exact age of the adult geese we marked, we knew they had nested as indicated by an incubation patch and cloacal characters (Hanson, 1959, 1965). All marked geese that did not attempt to nest were paired when they returned to the study area. We could not determine, however, why only 70% attempted to nest. Moser and Rusch (1989) reported nesting rates for interior Canada geese (Branta canadensis interior) approaching 100% for 5-7 year-old females and Brakhage (1965) found 84-87% of giant Canada geese older than 4 y old nested. Failure of some adult female geese to nest (Schmutz and Morse, 2000) and other negative effects of neck collars on geese (Ankney, 1975; Craven, 1979; Zicus et al., 1983; MacInnes and Dunn, 1988; Schmutz et al., 1997) have been reported. In this study, it is possible that radio collars had a detrimental effect on nesting attempts by geese. If that is the case, it is logical to conclude that the 70% nesting rate was a minimum value for this population.
Ground nests can be less successful than other nest types (Brakhage, 1965; Cooper, 1978; Perkins and Klimstra, 1984; Coluccy, 2001). Although only nine geese nested on shorelines during our study, 78% of those nests hatched. Muskrat mounds are important for Canada goose nesting (Cooper, 1978) and geese often used small clumps of cattails and other vegetation where there were no muskrat mounds. On these nests, flooding was the main cause of nest failure. Both apparent and Mayfield estimates of nest success indicate that reproductive success of giant Canada geese is excellent in eastern South Dakota, although predation and flooding suppress production in some areas. Egg success and hatching success were comparable to estimates reported for other populations of giant Canada geese (Table 3).
Mean TCS and GLN were higher than estimates during the reintroduction period in South Dakota (Table 3). In addition, clutch sizes of Canada goose nests were similar on all nest types, which is not always the case (Brakhage, 1965; Hilley, 1976). We did not observe a linear decline between clutch size and time of nest establishment which has been reported in other studies (Brakhage, 1965; Johnson, 1977; Sayler, 1977; Cooper, 1978; Nigus and Dinsmore, 1980; Coluccy, 2001).
The distances from goose nesting locations to wetlands where they molt are not well documented. Zicus (1981) reported the distances between nesting locations and wetlands used for brood rearing ranged from 0.7-8.4 km in Wisconsin, with 27% of families moving more than 7.5 km. Geese in South Dakota did not move to this extent, probably due to the abundance of wetlands and brood rearing habitats near nesting sites. Geese used permanent wetlands for brood rearing and molting, but not for nesting unless islands were available. Geese often nested on small wetlands that had stock dam berms, cattails mounds, and islands and then moved to larger wetlands to molt and raise their brood. Permanent wetlands are not used much for nest sites in eastern South Dakota, especially during wet years. Geese prefer the numerous small wetlands with ideal nesting sites such as muskrat huts (Mammenga 2002).
Geese that nested on the wetland where we captured them remained on that wetland throughout the brood rearing and molting period, which has been noted at other sites (Giroux, 1980; Zicus, 1981). We believe that these established rearing areas were where the adults had molted the previous summer. However, many geese moved their brood from the wetland where they nested to other wetlands for molting. After the brood reached the marked female's capture/molt wetland, no more brood movements to other wetlands occurred. Geese in South Dakota appear to use the same brood-rearing/molting wetlands year after year, as similarly observed in other studies (Zicus, 1981; Eberhardt et al., 1989; Didiuk and Rusch, 1998; Coluccy, 2001). Many geese moved their broods considerable distances through several wetlands to brood rearing wetlands indicating geese may be enticed to move to specific brood rearing sties. This is an important consideration for waterfowl biologists when they are making management decisions related to giant Canada geese.
Acknowledgements.--This work was supported by the South Dakota Department of Game, Fish and Parks, South Dakota State University, and the Federal Aid to Wildlife Restoration Fund (Project W-75-R, No. 7598) administered by the South Dakota Department of Game, Fish and Parks. We thank L. Flake, P. Mammenga and the referees for constructive comments on the manuscript.
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CHARLES D. DIETER (2) AND BOBBY J. ANDERSON (1) Department of Biology/Microbiology South Dakota State University, Brookings 57007
(1) Assistant Professor, Department of Biology, Valley City State University, Valley City, North Dakota 58072
(2) Corresponding author: e-mail: firstname.lastname@example.org
TABLE 1.--Proportion of VHF marked female Canada geese within each reproductive status in eastern South Dakota, 2001-2004 Successful Unsuccessful Nonbreeding Year n (%) n (%) n (%) 2001 15 (60.0) 5 (20.0) 5 (20.0) 2002 14 (50.0) 5 (17.9) 9 (32.1) 2003 12 (44.4) 5 (18.5) 10 (37.0) 2004 3 (37.5) 3 (37.5) 2 (25.0) Total (n = 88) 44 (50.0) 18 (20.5) 26 (29.5) TABLE 2.--Mayfield and apparent nesting success for VHF marked Canada geese in eastern South Dakota, 2001-2004 Nest Success (%) Year Nests n Mayfield Method Apparent Method 2001 20 (65.5) (75.0) 2002 19 (67.3) (73.7) 2003 17 (61.9) (70.6) 2004 6 (45.8) (50.0) Total 62 (62.7) (71.0) TABLE 3.--Selected reproductive parameters for several populations of giant Canada geese Location TCS GLN Nest Success % Eastern South Dakota 5.73 5.02 71.0 (A) 62.7 (M) Western South Dakota 5.30 4.60 78 (A) Western South Dakota 5.27 -- 57 (A) Northeastern South Dakota 5.20 4.70 87 (A) Northeastern South Dakota 5.44 -- 95 (A) Western South Dakota 4.90 4.63 78 (A) Northwest Iowa 6.00 5.19 79 (A) Northcentral Iowa 5.30 -- -- Barrington, Illinois 5.00 4.22 57 (A) Dog Lake, Manitoba 5.11 5.15 46 (A) Marshy Point, Manitoba 5.60 5.45 75 (A) Southeast Michigan 5.40 4.40 82 (A) Twin Cities, Minnesota 5.60 4.80 68 (A) Central Missouri 6.05 5.43 75 (M) North Dakota 5.85 5.06 69 (A) Pennsylvania 5.17 -- 73.5 (A) 60.0 (M) Nesting Hatch Egg Season Location Success % Success % Length Eastern South Dakota 89 63 58 Western South Dakota 68 -- 71 Western South Dakota -- 59 71 Northeastern South Dakota -- 78 ~57 Northeastern South Dakota 88 -- -- Western South Dakota -- 72 76 Northwest Iowa 86 77 78 Northcentral Iowa 83 52 -- Barrington, Illinois 79 58 74 Dog Lake, Manitoba 96 51 61 Marshy Point, Manitoba 97 67 70 Southeast Michigan 91 70 -- Twin Cities, Minnesota 96 61 -- Central Missouri 88 76 99 North Dakota 95 64 -- Pennsylvania 83 64 64 Location Source Eastern South Dakota This study Western South Dakota Lengkeek (1973) Western South Dakota Bultsma (1976) Northeastern South Dakota Hilley (1976) Northeastern South Dakota Clausing (1979) Western South Dakota Stiefel (1980) Northwest Iowa Nigus and Dinsmore (1980) Northcentral Iowa Zenner and LaGrange (1998) Barrington, Illinois Kossack (1950) Dog Lake, Manitoba Klopman (1958) Marshy Point, Manitoba Cooper (1978) Southeast Michigan Kaminski et al., (1979) Twin Cities, Minnesota Sayler (1977) Central Missouri Coluccy (2001) North Dakota Johnson (1977) Pennsylvania Jacobs and Dunn (2004) Apparent (A) nesting success Mayfield (M) nesting success
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|Author:||Dieter, Charles D.; Anderson, Bobby J.|
|Publication:||The American Midland Naturalist|
|Date:||Oct 1, 2009|
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