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Population Dynamics of Breeding Waterfowl

VII. Composite Reproductive Statistics


The number of young birds produced can be expressed as the product of the components just discussed: the number of potential breeding birds, proportion of these that breed, clutch size, nest success, and brood survival. The product of the last four components yields the number of young produced per bird, a measure of reproductive rate. An understanding of the dynamics of production is best gained by separately examining each component. Nonetheless, valid inferences about sources of variation in overall reproductive performance can be based on estimates or indices of reproductive rate. Here we briefly discuss some indices of reproductive rate and review some investigations into sources of variation in them.

The number of young produced can be estimated for local areas by direct counts of broods (Chapter 13). An index to duck production on important North American breeding areas is also obtained on a much larger scale by aerial surveys conducted each July (Henny, Anderson, and Pospahala 1972). These estimates of numbers of young are sometimes translated into brood-pair ratios, which can be thought of as estimates of reproductive rate. The translation generally uses estimates of numbers of adults or pairs on an area at the beginning of the breeding season (see Chapter 13).

Estimates of age ratios (young per adult or young per adult female) at certain times of the year also provide useful indices of reproductive rate. For several species of geese it is possible to distinguish visually young and adult birds at a distance, either on migration stopover areas or on wintering grounds, to estimate age ratio (Hewitt 1950, Boyd 1959, Lynch and Singleton 1964, Jones 1970). For most ducks, young cannot be distinguished from adults in the field, but age ratios among harvested birds are sometimes used to index reproductive rate (Petrides 1949, Bellrose et al. 1961). If adult and young birds available in a population during a hunting season do not have equal chances of being harvested, the age ratio in a harvest sample will provide a biased estimate of the population age ratio. It is often possible, however, to estimate these different probabilities of being harvested independently with band recovery data and to use such estimates to "correct" or adjust harvest age ratio data to yield an estimate of the age ratio in the population at the time of banding (Bellrose et al. 1961, Munro and Kimball 1982).

In addition to these indices of reproductive rate, detailed investigations using marked birds permit direct estimates. Such studies have also yielded inferences about sources of variation in reproductive rate.

1. Species and Geographic Variation

Several investigators have used brood-pair ratios to compare reproductive performance of species on local study areas (Table 14-3), although differential brood visibility may influence comparisons (Rumble and Flake 1982).

Table 14-3. Observed brood-pair ratios of North American waterfowl
Species
Area
Ratio
Source
Canada Goose Ont. 0.47 Dennis & North 1984
Wood Duck Ont. 0.38 Dennis & North 1984
American Wigeon Alb.
Sask.
0.39
0.30
Smith 1971
Leitch & Kaminski 1985
Gadwall Sask. 0.22 Leitch & Kaminski 1985
Mallard Alb.
Ont.
Sask.
Sask.
0.31
0.76
0.31
0.32
Smith 1971
Dennis & North 1984
Leitch & Kaminski 1985
Stoudt 1971
Northern Pintail Sask.
Sask.
0.35
0.20a
Leitch & Kaminski 1985
Stoudt 1971
Blue-winged Teal Alb.
Ont.
Sask.
Sask
0.41
0.43
0.43
0.31
Smith 1971
Dennis & North 1984
Leitch & Kaminski 1985
Stoudt 1971
Northern Shoveler Sask. 0.37 Leicht & Kaminski 1985
Canvasback Alb.
Sask.
0.62
0.62
Smith 1971
Stoudt 1971
Lesser Scaup Sask. 0.22 Leicht & Kaminski 1985
aPossibly biased low because pairs may have been counted on study area but moved elsewhere to nest (Stoudt 1971).

Bellrose et al. (1961:466) adjusted Mississippi Flyway harvest age ratios for differential vulnerability and compared the resulting age ratio estimates (juveniles per adult hen) for 12 species of ducks during 1946-49. Relatively high age ratios were calculated for Mallards, Black Ducks, Northern Shovelers, Ring-necked Ducks, and Canvasbacks. American Wigeon and Green-winged Teal exhibited intermediate age ratios. Age ratios were low for Gadwalls, Pintails, Blue-winged Teal, Redheads and Lesser Scaup. In a comparison of harvest age ratio data from southwestern Ontario (uncorrected for differential vulnerability), Dennis, Fischer, and McCullough (1984) found that Mallard age ratios were consistently higher than those of Black Ducks, suggesting greater Mallard production.

Pospahala, Anderson, and Henny (1974) found that the average number of duck broods (all species combined) per square mile and average number of broods per July pond exhibited a decreasing cline from west to east across the prairie-parkland region of Canada. Smith (1970) used aerial breeding ground survey data and harvest age ratio information to infer that Pintail reproductive rate was lower in northern than in southern breeding areas, a conclusion later corroborated by Calverley and Boag (1977). Using pair and brood counts, Dzubin and Gollop (1972) reported the proportion of hens fledging broods in a Manitoba parkland (0.40) to be nearly twice as large as that in a Saskatchewan prairie (0.21).

2. Age, Breeding Experience, and Social Status

Boyd (1959) examined age ratios taken from direct counts of White-fronted Geese made during December in England for four consecutive years. The temporal sequence of high and low age ratios led Boyd (1959) to conclude that the age structure of a population can be an important determinant of reproductive rate. Trauger (1971) found an increase in the brood-hen ratios of Lesser Scaup from 4.8% among yearlings to 47.4% among three- or four-year-olds. Ratcliffe, Rockwell, and Cooke (1988) calculated the number of young from female Snow Geese recruited to the breeding population and found that the recruitment rate increased with age of female until five to seven years, and declined for older ages.

Teunissen, Spaans, and Drent (1985) noted that for Brant competitive status affected reproductive success. For four pairs observed in several years, the highest ranking male raised the most young.

3. Wetland Habitat

Wetland habitat conditions on the breeding grounds are believed to influence several components of reproduction, as discussed previously. These separate effects combine to produce an important effect on overall production and reproductive rate. Harvest age ratio data (either corrected or uncorrected for differential vulnerability) have been used to investigate the relationship between Mallard reproductive rate (at both the continental and flyway levels) and May and July pond numbers on important North American breeding areas. Crissey (1969) found a strong correlation between the estimated continental production of Mallards and the number of July ponds in prairie Canada. Others have since used these same data in conjunction with both linear (Geis, Martinson, and Anderson 1969; Martin, Pospahala, and Nichols 1979; and nonlinear (Hammack and Brown 1974; Anderson 1975a, b; Brown, Hammack, and Tillman 1976) models and obtained strong positive associations between pond numbers and numbers of young Mallards produced. Predictions based on these models have been fairly good and have been used during the process of setting annual hunting regulations (Martin, Pospahala, and Nichols 1979).

Bellrose et al. (1961) plotted the number of May ponds per square mile in Manitoba and Saskatchewan, 1948-59 and the Mallard age ratio (young per adult) in hunter bag checks in the Mississippi Flyway. Ponds and age ratios covaried directly. Heitmeyer and Fredrickson (1981) and Kaminski and Gluesing (1987) used linear regression to investigate the relation of age ratio of Mallards in the Mississippi Flyway harvest in relation to spring population size, spring wetland conditions, and winter wetland conditions. In each analysis, age ratios were positively related to both spring and winter wetland indices.

Other workers have used pair and brood count data in conjunction with wetland counts on local study areas. Smith (1971) and Stoudt (1971) computed brood-pair ratios on their study areas for three time periods: predrought years (1953-58), drought years (1959-63), and postdrought years (1964-65). On the Lousana, Alberta study area, brood-pair ratios were lower during the drought period than during the other two periods for all four species studied: American Wigeon, Mallard, Blue-winged Teal, and Lesser Scaup (Smith 1971). On the Redvers, Saskatchewan study area, however, brood-pair ratios were not reduced during the drought years for Mallard, Pintail, Blue-winged Teal, or Canvasback (Stoudt 1971). Stoudt (1971) concluded that the Redvers area was one of the better waterfowl production areas in Canada, producing ducks even when most other areas were too dry to do so. Kaminski and Gluesing (1987) used brood-pair ratios for Mallards at Redvers during 1952-77 and found no evidence of a relation between this ratio and either May or August pond numbers.

Leitch and Kaminski (1985) computed brood-pair ratios for seven duck species on a Caron, Saskatchewan study area and tested for associations between these ratios and both May and August wetland numbers for 1950-75. Significant positive associations were found with August, but not May, pond numbers for all species (American Wigeon, Gadwall, Mallard, Pintail, Blue-winged Teal, Northern Shoveler, and Lesser Scaup).

4. Weather

Weather has direct effects on reproduction in addition to those involving wetlands. Bellrose et al. (1961) noted that much of the year-to-year variation in the age ratio of Mallards from Mississippi Flyway hunter bag checks could be explained by variation in May pond numbers for prairie Canada. For several of the years with lower age ratios than expected based on pond numbers alone, spring temperatures were well below normal, and spring blizzards occurred in some cases. Bellrose et al. (1961) concluded that cold and excessively wet springs may be as unfavorable to duck production as dry and warm springs.

Hammond and Johnson (1984) investigated the influence of weather on duck production using brood-pair ratios for two North Dakota study areas. For one area, but not the other, they found a significant positive relationship between the brood-pair ratio and mean temperatures between 23 April and 3 June for Mallard, Blue-winged Teal, and Redhead.

Stoudt (1982) mentioned cold spring temperatures, as well as flooding caused by excessive spring rainfall, as weather factors reducing Canvasback brood-pair ratios during some years. Sleet and cold during the summer were reported to be the "worst single hazard" to Snow Geese breeding at James Bay, and reduced age ratios were found for two years of "very severe" weather on the breeding grounds (Hewitt 1950).

Reeves, Cooch, and Munro (1976) found with remotely sensed imagery that snow and ice conditions in June on the breeding grounds of several species of arctic-nesting geese were related to age ratios in the subsequent harvest.

5. Density of Birds

From an analysis of aerial survey data (May counts of adult birds, July counts of broods) for all ducks combined, Pospahala, Anderson, and Henny (1974: 43-45) found no evidence that reproductive rate varied as a function of breeding population size. Kaminski and Gluesing (1987) used harvest age ratios of Mallards in the Mississippi Flyway to address the question of density-dependent reproductive rates. They used May population size estimates and winter survey counts as indicators of population size. Age ratios were found to be negatively related to both spring and winter population indices. Spring population size was also found to be a significant contributor to variation in age ratio when spring and winter wetland conditions were included in a multiple regression model. Similar analyses were conducted for subsets of the data corresponding to years when spring wetland conditions were wet, dry, and average. Age ratio was found to be negatively related to spring population size in wet, but not in dry or average years.

Density-dependent influences on reproductive rate have also been investigated using long-term data on brood and pair numbers for local study areas. Pospahala, Anderson, and Henny (1974) found linear relationships between duck brood and pair numbers for study areas in Alberta and Saskatchewan, with no indication of the nonlinearity indicative of density-dependent reproduction. Kaminski and Gluesing (1987) used different methods with Mallard data from two Saskatchewan study areas for 1950-75. They found that the brood-pair ratio on one study area, but not the other, was inversely related to the continental wintering population counts.

Ebbinge (1985) detected decreased proportions of successful pairs and brood size among Bean Geese and Brant in years with higher population densities. Patterson, Makepeace, and Williams (1983) noticed reduced age ratios of Northern Shelducks with higher densities of territorial pairs on an estuary.

6. Synopsis

Composite reproductive measures, including brood-pair ratios and age ratios, exhibit relationships similar to those involving the individual components of reproductive rate. Ratios tend to be higher in years with good wetland conditions and lower in years with adverse weather or high densities of breeding birds.
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