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Population Dynamics of Breeding Waterfowl

III. Proportion of Birds that Attempt to Breed

B. Timing of Breeding


The timing of waterfowl breeding seasons probably evolved in response to a variety of potential selective pressures. Day length and temperature are major proximate determinants of breeding (Lack 1954), mediated by hormones. We concentrate on other proximate factors.

A longer span of breeding allows for more nesting attempts and potentially greater reproduction. In some instances earlier breeding is advantageous for selection of a nest site (Grice and Rogers 1965, Findlay and Cooke 1982).

1. Species and Geographic Variation

Most waterfowl in temperate North America breed in spring or early summer. In temperate South America, breeding can occur in spring, fall, or both seasons, and tropical species may show less obvious patterns (Weller 1975). Because environmental constraints on breeding vary geographically, comparisons among species should be made within an area, as was done by Sowls (1955); Vaisanen (1974); Newton and Campbell (1975); Toft, Trauger, and Murdy (1982); Duebbert and Frank (1984); and Hammond and Johnson (1984). Tome (1984) proposed that, as a species, Ruddy Ducks do not breed soon after arrival on their breeding grounds because they are physically unable to accumulate necessary nutrient reserves except on the breeding ground.

Within a species, the timing of breeding often varies geographically. The onset of breeding is less fixed, and the duration often longer, in areas with favorable climate (reviewed by Newton 1977). Southern-breeding populations of a species typically initiate nesting earlier and continue longer than their northern-breeding counterparts. This holds for the Canada Goose (Klopman 1958, Mickelson 1973), Wood Duck (Cunningham 1968, Haramis 1975), Black Duck (Reed 1970), Mallard and Pintail (Calverley and Boag 1977), captive Mallard (Batt and Prince 1978), and several dabbling and diving duck species (Toft, Trauger, and Murdy 1982). Vaisanen (1974) found that nesting by early breeders, but not late breeders, was progressively delayed the farther north the breeding area. In addition, breeding may be later at higher altitudes (Krohn and Bizeau 1980 for Canada Geese).

2. Age, Breeding Experience, and Condition

Young or inexperienced hens begin nesting later than older or experienced ones in a variety of species: Mute Swan (Birkhead, Bacon, and Walter 1983), captive Hawaiian Goose (Kear 1973), Canada Goose (Brakhage 1965, Akesson and Raveling 1984), Wood Duck (Grice and Rogers 1965, Baker 1971b), Gadwall (Gates 1962, Blohm 1979), Mallard (Dzubin 1969, Krapu and Doty 1979), Black Duck (Coulter and Miller 1968, Reed 1970), Pintail (Duncan 1987a), Blue-winged Teal (Dane 1965), Northern Shoveler (Mednis 1968), Canvasback (Olson 1964), Redhead (Johnson 1978, Alliston 1979a), Tufted Duck (Mednis 1968), Eider (Korschgen 1976, Gerell 1985), and Goldeneye (Dow and Fredga 1984). In a few species, nest initiation date appears to differ among more than two age classes: Snow Goose (Finney and Cooke 1978, Hamann and Cooke 1987), Gadwall (Lokemoen, Duebbert, and Sharp 1990), Lesser Scaup (Afton 1984), and Goldeneye (Dow and Fredga 1984).

Despite starting earlier, older birds may persist with laying attempts longer than younger ones (Olson 1964 for Canvasback; Lokemoen, Duebbert, and Sharp [1990] for Gadwall). The earlier and more prolonged nesting by older, more experienced waterfowl has been variously attributed to their better condition (Mihelsons et al. 1968, Ely and Raveling 1984), greater food-gathering abilities (Duncan 1987a), earlier arrival on breeding grounds (Johnson 1978), and superior home ranges (Dzubin 1969).

There is both direct and indirect evidence that condition of the female influences the timing of nesting. Earlier laying by females in better condition was indicated by Spurr and Milne (1976) for Eiders, J. T. Barnett and J. K. Ringelman (pers. commun.) for Mallards, and Hepp et al. (1987) for Wood Ducks. Captive Mallards on restricted diets initiated laying later than those on unrestricted diets (Pattenden and Boag 1989). Indirect evidence that condition affects timing of nesting includes late nesting the following year by Canada Geese that had had larger broods (Lessells 1986), late laying dates of Mute Swans following a severe winter (Bacon 1980), and late breeding by Black Swans after a winter of reduced food supplies (Miers and Williams 1969).

3. Wetland and Nesting Habitat

The effect of wetlands as feeding and resting sites on the timing of waterfowl breeding is readily apparent. On North American prairies, delayed nesting or abbreviated nesting seasons can result from drought (Rogers 1964, Afton 1984). Gauthier (1987a) noticed earlier laying by Buffleheads on territories with greater food densities. In northern areas, the breakup of ice on the feeding grounds affects the timing of waterfowl nesting (Belopol'skii 1961). In other areas as well, water levels, especially as they affect food supplies, may influence the timing of breeding seasons: Laysan Duck in Hawaii (Moulton and Weller 1984), Ruddy Duck in California (Gray 1980), and various waterfowl species in Australia (Braithwaite and Frith 1969). In interior Australia, where rainfall is erratic, direct factors, such as food supply, and factors predictive of foods, such as rainfall or water levels, affect breeding (Braithwaite 1976). Black Swans bred regularly in some parts of their range, but in other parts they bred opportunistically in response to wetland availability (Braithwaite 1970).

The availability of nest sites can influence timing of breeding. Nesting on islands is often deferred until they are no longer ice-bound (Hildén 1964, Ahlén and Åndersson 1970), probably as a safeguard against terrestrial predators (Lack 1950, Schamel 1977). Nest sites on upland areas must be free of snow, ice, and meltwater before nesting commences (Barry 1967, Ryder 1967, Mednis 1968, Eisenhauer and Kirkpatrick 1977, Kerbes 1986). Waterfowl that nest in vegetation may delay breeding until vegetation reaches an appropriate stage for nest construction or security (Mendall 1958, Meanley and Meanley 1959, Havlín 1966b, Jarvis and Harris 1971).

4. Weather

Weather affects the timing of the nesting season in several manners, in addition to the effects mediated by availability of food and nesting habitat. Effects on the onset of breeding were recently reviewed by Hammond and Johnson (1984). Associations between weather factors and the onset of breeding generally involve delays caused by cold temperatures or severe storms, but other weather phenomena have also been implicated, including precipitation (Mendall 1958) and wind (Ryder 1967).

Early-nesting species are more often affected by weather than late-nesting ones (Mendall 1958, Bezzel 1962, Newton and Campbell 1975). Conversely, Hammond and Johnson (1984) showed that the later-nesting Gadwall and Blue-winged Teal were affected as much as the Mallard by cool temperatures during their normal initiation period. The difference between early- and late-nesting species may simply reflect the greater chance that early-nesting species will encounter adverse weather during nest initiation.

Nesting may be prolonged into the summer by certain weather features, such as adequate precipitation (Hammond and Johnson 1984 for Mallard and Blue-winged Teal, Lavery 1967 for Wandering Whistling Duck and Grey Duck), and by above-normal temperatures in spring (Hanson and Eberhardt 1971 for Canada Goose). Delayed start of breeding due to inclement spring weather may result in an extended nesting period (Sowls 1955 for Mallards and Pintails; Cowardin, Gilmer, and Shaiffer 1985 for Mallards).

5. Other Factors

Intraspecific competition can affect laying date; Blohm (1979) found earlier and more synchronous nesting in Gadwalls during years of lower breeding density. Several investigators have found that individual females exhibit high year-to-year repeatability of laying date: Bacon (1980) for Mute Swan, Cooke and Rockwell (1988) for Snow Goose, Batt and Prince (1979) for captive Mallard, Spurr and Milne (1976) and Laurila and Hario (1988) for Eider, Koskimies (1957) for White-winged Scoter, and Dow and Fredga (1984) for Goldeneye.

Some social factors have been proposed as influences on the timing of breeding: time of pairing the previous winter for Eiders (Spurr and Milne 1976), and social stimulation for Eurasian Pochards (Bezzel 1969).


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