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Breeding Population Inventories and Measures of Recruitment

II. Population Estimates

B. Obtaining the Count


A host of methods has been used for obtaining counts of breeding waterfowl. These vary depending on the purpose for which the data are to be used, the size of the study area or sample plot, and the species to be inventoried.

1. Ground Counts

Ground counts of waterfowl are used for detailed local studies as well as on sample plots from which estimates are extrapolated to extensive areas. They have the advantage of allowing the type of biological interpretation discussed in the previous section. Such interpretation is essential to the accuracy of estimates on local areas. The disadvantage of ground counts is their cost in terms of labor and money and access to waterfowl breeding areas, especially for geese, which often breed in remote or inaccessible areas. Stott and Olson (1972) evaluated both ground counts and aerial surveys for sea ducks on the Atlantic coast and concluded that, because of variability in aerial counts, ground counts were superior.

Many species of waterfowl frequent areas where they are hidden by vegetation. Biologists have coined the term "beat out" to describe a survey in which birds are flushed from cover in order to tally them. This method involves walking through all areas of vegetation and often includes making noise to assist in flushing the birds. On deeper water areas similar techniques have been used from canoes (Stoudt 1938, Collins 1974). Besides being an arduous task, ground counts lead to another problem, for which the term "roll up" has been used. Birds flushed from one wetland may settle on another, and the same birds may be counted a second time. In addition, the act of flushing may make the behavioral interpretations discussed in the previous section difficult.

Counting birds from stationary observation points and assuming that all birds will move into openings and become visible is an alternative to "beat outs" (Kadlec 1966). Rumble and Flake (1982) compared active and passive counting techniques for broods, which are more difficult to observe than pairs. Kirby (1980) used passive counts from elevated blinds for both pairs and broods in forested areas of Minnesota where direct counts were difficult or impossible. In some habitats passive methods will not work because birds may never move into areas where they are observable. Passive counts also present sampling problems because of the difficulty in determining whether birds counted from one observation point are the same birds as those counted from another. For prairie habitat, Dzubin (1969a) recommended a combination of methods that included counting pairs from a vehicle parked at a good observation point for each pond and using noise to flush birds not seen from the observation point.

There can be no hard-and-fast rules about the use of active flushing versus passive counting from observation points. The method must vary depending on the nature of the habitat, the size of the wetlands, and the ease of access to them. Under some habitat conditions a direct method may be virtually impossible, especially for broods, and one of the less direct methods listed below must be used.

2. Counts from Aircraft

The inaccessibility of waterfowl habitat and the vast areas over which waterfowl breed led to the use of aircraft for making inventories. As early as 1947, Sowls (1947:449) stated, "They [USFWS] have added a first-class technician to the Mississippi Flyway and have made full use of aircraft which we now know to be indispensable in the breeding grounds." In fact, inventory of waterfowl has led the way in the use of aircraft in wildlife management. The cooperative breeding ground surveys (Martin et al. 1979) conducted in North America are totally dependent on the use of aircraft. Aerial surveys for estimating waterfowl numbers are far more extensive than the area surveyed by federal agencies in the United States and Canada. Similar techniques have been used by states including Minnesota (Jessen 1970), Wisconsin (March et al. 1973), and Colorado (Kinghorn 1949) as well as by the provinces in eastern Canada (Gillespie and Wetmore 1974, Kaczynski and Chamberlain 1968). Aerial surveys over large areas have also been used in northern Fennoscandia (Haapanen and Nilsson 1979), Estonia (Lilleleht 1976), Australia (Braithwaite et al. 1986), and France (Dervieux et al. 1980).

Aerial surveys for ducks have generally been conducted from fixed-wing aircraft flying at low level, but helicopters have also been used especially for arctic-nesting geese (Reed and Changnon 1987). Helicopters have advantages over fixed-wing aircraft because they can be flown at low speeds and low elevations, and they furnish excellent visibility. The noise from a helicopter may disturb birds and move them. The cost of helicopters is much higher than for fixed-wing aircraft, and they have not been used as the primary tool in continental surveys. Ross (1985) found that helicopter and ground surveys for conducting waterfowl counts in the boreal forest had comparable costs when salaries were considered. In addition, he found that the Black Ducks moved into open water where they could be counted when approached by a helicopter. Kaminski and Prince (1984) used a helicopter to count pairs in marsh habitat and found that 50 64.8-ha plots could be censused in four hours.

Conspicuous species such as swans can be inventoried from aircraft with reasonable certainty that all individuals will be counted (King 1973, Lensink 1973, King and Conant 1981, Reese 1982). Other species are difficult to observe or to identify, especially females (McLaren and McLaren 1982). Aircraft counts generally miss some individuals for most species of waterfowl even when the survey is designed to obtain a total count. Furthermore, the proportion of the population that is missed will vary with species, habitat, year, and to some extent the observer (Diem and Lu 1960, Tacha et al. 1978, Broome 1985). Caughley (1974) discussed various biases that can occur from aircraft transect censuses of big game, and Caughley and Goddard (1972) suggested a method for estimating the number of animals missed by using repeated censuses, but Routledge (1981) presented evidence that their method was not appropriate. Burnham and Anderson (1984) discussed methods of estimating numbers from strip transect data where the distance to each animal observed is known. If the proportion of birds not seen remains constant, an assumption that is open to question, uncorrected aerial counts can serve as an index to population size. Normally, however, interest is in an estimate of size of the population.

One source of bias in aerial waterfowl inventories can be overcome by estimating the proportion of birds that are not seen by comparing birds seen from the aircraft with birds seen on the ground, which provides an airground correction ratio (U.S. Fish and Wildlife Service and Canadian Wildlife Service 1987). Nearly simultaneous counts of sample areas are made from an aircraft and by ground surveys. An air-ground correction ratio is then used to expand aerial counts by species on areas of similar habitat. These ratios are calculated each year during the cooperative breeding ground surveys in North America on those transects in the prairie habitat.

On northern transects in the forested region, constant ratios are used. Although many species are found here, they often occur in low densities, and the logistic problems and associated expenses in labor make extensive ground verification of aerial counts unfeasible. Consequently, aerial counts by species have been adjusted annually with long-term constants derived principally from historical data gathered in prairie-parkland areas. However, the necessity of obtaining annual adjustments in the boreal region has long been recognized. Beginning with preliminary trials in northern Manitoba and northern Saskatchewan in the late 1970s, and continuing with a more expanded effort across all northern areas in 1985 and 1986, helicopters are being used in place of ground counts to construct representative visibility rate estimates for the region.

The use of helicopters for censusing waterfowl has been considered for many years, and various studies have explored their feasibility in a variety of habitats and conditions (Lotter and Cornwell 1969, Bateman 1970, Malecki et al. 1981). The advantages of helicopter surveys include the speed with which the counts may be obtained, thus increasing sample size, and their potential use in remote areas where there is little chance of obtaining ground counts. However, the substitution of helicopters for ground counts also assumes that all birds are counted from the air, an assumption that may not be satisfied in many situations. In addition, helicopter flight time is expensive, which may limit its application. Ratios of helicopter to ground counts and of helicopter to fixed-wing counts have varied from study to study, but in some cases the results suggest that this survey method has promise.

Several assumptions are required for the corrected estimate to be without bias: (1) all individuals are counted in the ground count, (2) the population sampled from the aircraft is the same as the population sampled from the ground, and (3) the air-ground ratio derived from the sample plots is the same as that for unsampled plots to which it is applied. In practice, some of these assumptions may not be met. For example, the ground subsample of the cooperative breeding ground surveys is purposely selected and may not be representative of the aerial sample. In addition, ground access to some transects is not practical, and some of the usual assumptions regarding randomness and subsequent independence among the various estimators are not met. Nevertheless, some degree of bias is accepted and tempered by the high precision attained by very large sample sizes that are obtained with the aerial survey. Sorensen and Ibister (1970), Surrendi (1972), and Bowden (1974), among others, have discussed possible sources of bias associated with aerial surveys of ducks in prairie-parkland areas of Canada and the United States. Species identification from aircraft can also be a source of error in surveys, although experienced observers are very skillful in identifying birds from the air. However, in most aerial surveys of breeding pairs some birds will not be identified to species and are placed in an unidentified category. Species identification is even more difficult for broods, and in many surveys, including the cooperative breeding ground survey, no attempt is made to separate species.

3. Counts from Aerial Photographs

Aerial photographs can be used to obtain accurate counts of waterfowl. They have frequently been used to inventory birds in migration or in wintering flocks. More than 40 years ago, Spinner (1946) used aerial photographs to count Snow Geese in migrating flocks, and later Chattin (1952) used large-format aerial photographs of waterfowl concentrations in California. Chattin (1952) determined density of birds within flocks from sample plots on each photo but did not attempt to distinguish species. From high-quality photographs (e.g., Ferguson and Gilmer 1980), it is possible to identify species and sometimes sex (Ferguson et al. 1981). For some species, age can be determined (Heyland 1972, Reed and Changnon 1987). Provan (1976) used aerial photographs obtained along transect lines to estimate the number of ducks present and to calculate the variance of the estimate. Haramis et al. (1985) used photographs from a hand-held 35-mm camera to obtain sex ratio and flock size data for Canvasbacks in winter flocks. Mendall (1968) used aerial photographs to obtain supplementary sex ratio data for eiders. Vertical aerial photographs have been used frequently for determining the number of Snow Geese (Spinner 1946, Heyland 1972, Kerbes 1975, Bordage and Dupuis 1983). Heyland (1972) was also able to distinguish young from adults and to determine the size of family groups. McLandress (1979) determined the ratio of Ross' Geese to lesser snow geese by ground surveys in California and then applied that ratio to photographs of concentrations of white geese to estimate the size of Ross' Goose populations. Kerbes et al. (1983) used LANDSAT data to select islands where Ross' Geese nested and then photographed the nesting colonies. Ground-level photographs have been used to obtain indices to breeding waterfowl populations (Cowardin and Ashe 1965). This technique is suitable for comparison of habitat use among areas or time periods in research studies but is too expensive and complex to be used over large areas.

Erwin (1982) used aerial photographs experimentally in a test of the ability of different observers to count large numbers of ducks in flocks. He found that without reinforcement through training, experienced observers were more accurate than inexperienced observers. Eight of nine observers tested estimated number of birds within 10% of the actual number.

4. Indices to Breeding Population

Winter or migration counts have been used as an index to breeding population for species that cannot be inventoried on their breeding grounds. For example, the Black Duck has sparse breeding populations spread over a vast area of forested habitat, presenting almost insurmountable inventory problems. The breeding grounds of some species, such as the tule subspecies of White-fronted Geese, have only recently been discovered (Timm et al. 1982). Winter surveys have been used in North America since the mid 1930s. Prior to initiation of the cooperative breeding ground surveys, they provided the primary source of information on waterfowl abundance in North America (Martin et al. 1979). The accuracy of winter inventories is unknown and results obtained from them sometimes are not consistent with those obtained from extensive breeding ground surveys. Recently, because of concern over the status of the Black Duck, the USFWS initiated research to determine whether winter surveys for that species can be improved by using a statistically valid sampling design similar to that used in the breeding ground surveys (Conroy et al. 1988).

Extensive surveys by cooperating professional and amateur ornithologists are used for estimating waterfowl numbers in Europe (Milstein 1968, Yarker and Atkinson-Willes 1972). In North America a breeding bird survey is conducted by cooperating amateur ornithologists along roadside transects (Erskine 1978, Robbins et al. 1986). The surveys do not estimate the size of breeding populations but do yield an index to abundance that has been used to establish population trends for all species. A summary of population trends from the first 15 years of the survey has recently been published (Robbins et al. 1986).

Because of their selection of woodland habitats, Wood Ducks are one of the most difficult waterfowl species to inventory, and special survey techniques have been developed. Smith and Flake (1985) found that on a prairie river Wood Duck broods were restricted to the river and adjacent oxbow lakes. Several states have calculated population indices for Wood Ducks based on counts made from a boat floating stretches of river, but these indices are subject to many biases and lack precision (Cink 1977).

Postbreeding Wood Ducks use roosting areas at night and fly to them at dusk. Counts of birds entering these areas have been used as an index to Wood Duck abundance (Luckett and Hair 1979). Hein and Haugen (1966) concluded that roost counts were sufficiently precise to detect a 15% change in population size. The validity of the technique has been questioned because of daily (Tabberer et al. 1971) and yearly variation in use of roosts. Parr and Scott (1978) reviewed use of roost counts as a population index and concluded that the technique was not practical. Furthermore, roost counts, like winter counts, include both surviving adults from the breeding population and recruits from that year's production and can furnish only a rough index to breeding population.

5. Estimates Based on Habitat Availability

A correlation between spring breeding populations of waterfowl and the number of ponds present on a local breeding area has been known for many years (Dzubin 1969b:205). Crissey (1969) and later Johnson and Grier (1988) reviewed these correlations for data from the cooperative breeding ground surveys for North America. Pond data are easy to obtain over vast areas by means of modern remote sensing technology (Koeln et al. 1986). Cowardin et al. (1983) developed predictive equations for estimating duck breeding populations from data on the number, size, and class of wetland basins. This approach has considerable merit when the correlation between pond numbers and habitat is good, but it will not work when there are insufficient ducks to fill the habitat. Johnson (1986) demonstrated that the relation is best near the center of a duck's breeding range but that it has been deteriorating in recent years in North America.

6. Indirect Estimates Based on Mathematical Models

Pairs of some species such as forest-dwelling ducks and broods of most duck species are extremely difficult to survey. A number of innovative techniques have been developed. Bennett (1967) estimated the total number of broods on a marsh by repeated censuses in which he recorded the number of previously unsighted broods on each census. He fitted a logarithmic function to the data and then used the function to estimate the number of sightings required to see all broods. The method was most applicable for single isolated impoundments. Cook and Jacobson (1979) presented a method of correcting for visibility bias in aerial surveys. Their method was designed for aerial census of big game species and requires the use of primary and secondary observers who independently observe the same area. They presented models for estimating the number of animals present and the variances of the estimates.

Waterfowl have been banded in large numbers and furnish an opportunity for development of indirect population estimates based on band recovery and harvest data. Crissey (1957) pointed out that if the size of the harvest can be estimated from harvest surveys and the rate of harvest can be estimated from banding data, then the fall population producing the harvest can be estimated by harvest divided by harvest rate. If summer survival rate is also known, then calculation of the size of the breeding population that produced the harvest is possible. These techniques have been used to estimate North American populations of Mallards (Crissey 1957) and Wood Ducks (Geis 1966).

Mark-recapture models have also been used for estimating population parameters (Boyd 1956, Anderson and Sterling 1974), but as Martin et al. (1979) pointed out, these methods are more appropriate for species that breed in relatively discrete populations than for continental populations. Sulzbach and Cooke (1979) used mark-recapture methods to estimate population parameters, including population size, for a Lesser Snow Goose population. Coulson (1984) banded nesting Eiders and recaptured them in subsequent years. He then used Jolly-Seber models (Jolly 1965, Seber 1965) to estimate population size. Barratt (1966) suggested the possibility of banding Wood Ducks by night-lighting, recapturing them by drive-trapping, and then using mark-recapture methods to estimate population size. D. G. Raveling (pers. comm.) used the Petersen index to estimate breeding population by treating sightings of neckbanded geese as recaptures on the wintering ground.


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