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Nesting Ecology and Nesting Habitat Requirements of Ohio's Grassland-nesting Birds: A Literature Review

Results and Discussion

Ring-Necked Pheasant

Ring-necked pheasants were introduced in Ohio in 1896 and became established in 10 counties by 1903. Between 1923 and 1935, 10,000-25,000 ring-necked pheasants were released annually. Ring-necked pheasant populations in Ohio peaked during the late 1930s and early 1940s. Despite annual releases, ring-necked pheasant populations began declining during the mid-1940s. Changing agricultural land-use practices precipitated the initial population decline and continue to dictate ring-necked pheasant population levels in Ohio today (Peterjohn 1989).

In Ohio, ring-necked pheasants may lay their first clutches in late April, but most are produced during the last half of May. The peak hatch normally occurs in June and young pheasants remain with the hen into August. Young pheasants usually become independent by early August, but renesting attempts result in hens being accompanied by partially grown young into the first half of September (Peterjohn 1989, Peterjohn and Rice 1991).

Being year-round residents, deficiencies in either nest-brood habitat or winter cover/food can limit ring-necked pheasant populations. A shortage of quality nesting and brood-rearing cover was considered the major limiting factor of ring-necked pheasant populations in the Midwest (Warner 1979, 1984; Warner et al. 1984; Warner and Etter 1985, 1986). Nesting and brood-rearing habitats are considered together as ring-necked pheasant broods >/=3 weeks old centered their activities within 2-12 ha of the nest site (Kuck et al.1970, Warner 1979). Habitat types commonly used by ring-necked pheasants for nesting and brood rearing included idle grasslands, hayfields, pastures, wetlands, woodlands, strip cover (roadside rights-of-way, fencerows, grassed waterways, railroad rights-of-way, and drainage ditch banks), small grains, cultivated rowcrops, and no-tillage corn and soybean fields (Baskett 1947; Leedy and Hendershot 1947; Leedy 1949; Salinger 1952; Klonglan 1955; Linder et al. 1960; Wright and Otte 1962; Joselyn and Warnock 1964; Warnock and Joselyn 1964; Gates and Ostrom 1966; Francis 1968; Joselyn et al. 1968; Bartmann 1969; Frank and Woehler 1969; Hanson 1970; Walcheck 1970; Erickson and Wiebe 1973; Gates and Hale 1974, 1975; Olson and Flake 1975; George et al. 1979; Braband 1984; Snyder 1984; Warner et al. 1984; Wooley et al. 1985; Basore et al. 1986; Warner and Joselyn 1986; Haensly et al. 1987; Meyers et al. 1988; Carroll and Sayler 1990; Lauber 1991; Warner 1992). In Ohio, Leedy and Hendershot (1947) estimated that 50-67% of all ring-necked pheasant nests were in hayfields. Gates and Hale (1974) considered wetlands and retired cropland the most important year-round cover types for ring-necked pheasants in Wisconsin. They reported that 63% of the annual pheasant production occurred in wetlands but noted that wetlands received little use by hens with broods.

Quality ring-necked pheasant brood-rearing habitat was described as having herbaceous cover that provided little obstruction to movement at ground level, good overhead protective cover, and abundant invertebrate populations (Warner 1979). Warner (1984) reported that ring-necked pheasant broods in Illinois used oats, hayfields, and cultivated rowcrops. Although oats and hayfields constituted only 6.4% of the study area, Warner (1979, 1984) noted that approximately half of all radiotelemetry locations for broods </=4 weeks old were in those 2 cover types. Either an oat field or a hayfield served as the primary focus of activity for all broods. Broods from hatch to 9 weeks of age covered an average of 17.8 ha in oat- and hayfield-dominated landscapes and 22.3 ha in rowcrop-dominated landscapes, suggesting that the former were of higher quality (Warner 1984). Corn and soybeans were considered of little value to ring-necked pheasant chicks as foraging habitat because of their low insect abundance and biomass (Warner 1979, 1984). In Minnesota, ring-necked pheasant broods used cool-season grass roadsides and warm-season grass plantings (Nelson et al. 1990). Cool-season grass roadsides were considered the better brood-rearing habitat because of their lower amount of litter (which impeded chick movement) and higher invertebrate abundance and biomass.

Several studies reported that ring-necked pheasants nested in herbaceous vegetation with an average canopy height of 18-25 cm, but vegetation at nest sites ranged in height from 25 to 69 cm (Hanson 1970, Gates and Hale 1975, Wood and Brotherson 1981). In Ohio CRP grasslands, ring-necked pheasant abundance was positively related to mean herbaceous canopy height (Swanson et al. 1995). Mean vegetation height-density at ring-necked pheasant nest sites ranged from 20 to 70 cm and herbaceous cover ranged from 48 to 68% (Wood and Brotherson 1981). Haensly et al. (1987) characterized vegetation at 181 ring-necked pheasant nest sites as 3-5% woody cover, 32-38% grass cover, 11-16% forb cover, 23-34% litter cover, and 19-29% bare ground. Because ring-necked pheasants prefer nesting in residual cover greater than or equal to 30 cm tall, a mean vegetation height-density greater than or equal to 20 cm was considered optimal for nesting and brood-rearing habitats whereas a vegetation height-density less than or equal to 5 cm was considered unsuitable (U.S. Fish and Wildl. Serv. 1987).

Ring-necked pheasants preferred herbaceous vegetation of medium height and density during early spring (pre-breeding season) and autumn (post-breeding season) (Pearce 1945, Gates and Hale 1974). Breeding territories of male ring-necked pheasants contained areas of tall, dense vegetation for loafing cover and open areas of short, sparse vegetation which permitted visual contact with other cocks, hens, and predators. Typical habitats occupied by crowing cocks included hayfields and lightly-grazed pastures intermixed with marshes and brushy vegetation (Baskett 1947, Burger 1966). During autumn, ring-necked pheasants avoided disturbed areas with short, sparse vegetation (e.g., heavily-grazed pastures and recently mowed hayfields) and selected idle grasslands and strip cover (Gates and Ostrom 1966). Corn fields, retired cropland planted to cool-season grasses, and marshes were important autumn habitats for radio-equipped ring-necked pheasant hens in Wisconsin (Gatti et al. 1989).

Several researchers considered brushy woodlands and marshes the most important winter cover types for ring-necked pheasants (Leedy and Hendershot 1947, Robertson 1958, Gates and Hale 1974, Sather-Blair and Linder 1980, Gatti et al. 1989, Leptich 1992). Woodlots, shelterbelts (i.e., narrow linear tracts of trees along field borders), and tall, persistent herbaceous vegetation, especially marshes dominated by bulrushes and cattails (Typha spp.), were used by ring-necked pheasants during winter as daytime roosting cover; low, open herbaceous vegetation (grasses, sedges, and grain stubble) was used for night roosting cover, even during severe winter weather (Robertson 1958, Skousen and Brotherson 1982, Warner and David 1982). Leedy and Hendershot (1947) considered woodlots with trees less than or equal to 10 m tall and shrubby understories valuable winter cover for ring-necked pheasants in Ohio. Ideal ring-necked pheasant winter habitat in Wisconsin was closed-canopy shrub-dominated wetland (Gates and Hale 1974). Shrubby areas with a canopy closure of greater than or equal to 30% were considered optimal winter cover for ring-necked pheasants (U.S. Fish and Wildl. Serv. 1987). On upland sites, Gates and Hale (1974) considered woody shelterbelts, especially those with coniferous trees, important ring-necked pheasant winter cover.

In Colorado, ring-necked pheasant night-time winter roost sites were characterized by vegetation greater than or equal to 38 cm tall and a plant density of 65-323 stems/m2 (Lyon 1954). Persistent herbaceous vegetation greater than or equal to 2 m tall provides optimal ring-necked pheasant winter cover; as mean height decreases below 1 m the quality of herbaceous vegetation as potential winter cover decreases (U.S. Fish and Wildl. Serv. 1987).

The annual diet of adult ring-necked pheasants consisted of 89% seeds, 5% plant foliage, 5% animal matter, and 1% mineral matter (grit) (Dalke 1937, Fried 1940, Korschgen 1964). Most seeds consumed by ring-necked pheasants were cultivated grains, corn being the most important (47-67% of the annual diet) (Korschgen 1964). Ring-necked pheasant hens increased consumption of insects and calcium-rich foods (snails) in spring, prior to egg laying (Fried 1940). Chicks fed predominantly on insects during the first 4-6 weeks posthatching, after which the amount of weed seeds and cultivated grains in the diet increased (Loughrey and Stinson 1955, Whitmore et al. 1986).

Corn was the primary winter food of ring-necked pheasants in most studies; however, other rowcrops may be regionally important. Major winter foods of ring-necked pheasants in Wisconsin were corn and oats, but burdock (Arctium spp.) seeds, nightshade (Solanum spp.) berries, and grassy plant material also were important (Gates and Hale 1974). As little as 15 cm of packed snow can cover waste grain in harvested fields and render it unavailable to ring-necked pheasants (U.S. Fish and Wildl. Serv. 1987). Thus, Gates and Hale (1974) considered unharvested corn, being tall enough to stand above 15 cm of snow, an important winter food source for ring-necked pheasants. Because ring-necked pheasants seldom foraged >0.4 km from cover during winter (Green 1938, Gates and Hale 1974), winter food and cover should be adjacent and well interspersed (U.S. Fish and Wildl. Serv. 1987).

Several studies reported that ring-necked pheasants moved up to 24 km between breeding and wintering areas (Sharp and McClure 1945, Kimball 1948, Warner and Etter 1986) but most reported movements </=3.2 km between these seasonal cover types (Green 1938, Robertson 1958, Hanson and Progulske 1973, Gates and Hale 1974, Lachlan and Bray 1976). In Wisconsin, Gates and Hale (1974) reported that >80% of marked ring-necked pheasant hens selected wintering areas within 3.2 km of their summer range and Dumke and Pils (1979) reported that all marked hens nested within 0.8 km of their winter range. Thus, ring-necked pheasant hens benefit most when nest-brood cover and winter cover are within </=3.2 km of each other.

The minimum area requirement of the ring-necked pheasant is unknown (U.S. Fish and Wildl. Serv. 1987). Gates and Hale (1974) suggested that year-round ring-necked pheasant management units be no smaller than 9 township sections (2,331 ha) centered on traditionally used winter cover. Gates and Hale (1974) recommended 8-12 ha of winter cover in each management unit. For maximum suitability, >/=0.5% of a management unit should provide ring-necked pheasant winter cover (U.S. Fish and Wildl. Serv. 1987). Recommendations for the amount of unharvested grain (corn or sorghum) </=0.4 km of winter cover in a management unit ranged from 0.2 to 1.2 ha (Frank and Woehler 1969, Gates and Hale 1974).

For maximum suitability, 30-50% of a ring-necked pheasant management unit should provide nesting and brood-rearing habitat (U.S. Fish and Wildl. Serv. 1987). Gates and Hale (1975) suggested that a minimum of 5% of a management unit be in nesting and brood-rearing habitat, but presumed that higher percentages would enhance reproductive success. In Illinois, Warner (1984) reported that the percentage of successful ring-necked pheasant nests annually from 1973 to 1981 was positively correlated with the amount of grassland habitat/hen in spring.

In Oregon, Haensly et al. (1987) reported that ring-necked pheasant nests in strip habitats (fencerows, roadside rightsof-way, and ditch banks) experienced rates of predation 4 times greater than nests in non-strip habitats (hayfields and pastures). They attributed the difference in predation rates to differences in habitat pattern (strip vs. non-strip) rather than vegetative structure at nest sites. Because ring-necked pheasant nest success was lower in strip than non-strip habitats in several studies (Eklund 1942, Baskett 1947, Gates et al. 1970, Gates and Hale 1975, Dumke and Pils 1979), blocks of undisturbed grass-legume cover >/=6 ha were recommended for optimal ring-necked pheasant nesting and brood-rearing habitat (Haensly et al. 1987).

Land use in most of the ring-necked pheasant's range is 50-95% cultivated cropland with 20-30% in corn and 20-40% in small grains (U.S. Fish and Wildl. Serv. 1987). In Illinois, Labisky et al. (1964) reported that ring-necked pheasant abundance was positively correlated with acreage in small grains and negatively correlated with acreage in pasture and woodland. Frank and Woehler (1969) reported that pheasant densities in Wisconsin were highest in townships with 55-70% of the area in cultivated cropland. They noted, however, that localized concentrations of ring-necked pheasants were associated with diverted cropland which provided undisturbed grass-legume cover.

Throughout the Midwest, ring-necked pheasant populations have decreased over the past 30 years coinciding with intensification of agricultural practices (Farris and Cole 1981, Dahlgren 1988). Shifts in production from hay and small grains to corn and soybeans reduced the amount of secure ring-necked pheasant nesting and brood-rearing habitat to critical levels (Warner 1979). Ring-necked pheasants readily nest in no-tillage rowcrops (Wooley et al. 1985, Basore et al. 1986). However, recent investigations indicated that widespread adoption of no-tillage planting methods in the Midwest will be of little value to ring-necked pheasants as reproductive success in these habitats is below replacement levels (Warner and Etter 1985, Wooley et al. 1985, Basore et al. 1986, Best 1986). The average number of ring-necked pheasant chicks hatched/successful nest in Illinois has remained relatively constant at about 8.6 since World War II (Warner et al. 1984). However, the average survival rate from hatch to 6 weeks has declined approximately 28% since the early 1950s (Warner et al. 1984). The loss of traditional brood-rearing habitat and the increased use of pesticides are considered the major factors affecting ring-necked pheasant chick survival. Hens with broods traditionally ranged close (</=400 m) to the site of hatch during the first 3-4 weeks posthatching, primarily in fields of oats and forage legumes (Warner 1979). Because small grains and hay have been replaced by rowcrops, broods now range over larger areas to meet their nutritional needs and have become more vulnerable to mortality by predation or separation from the hen (Warner 1984). Herbicides, by eliminating weedy habitats that support a diverse and abundant insect fauna, have reduced insect food availability for pheasant chicks (Warner et al. 1984).

The Conservation Reserve provision of the Soil Bank established perennial grasses and legumes on cropland during the 1960s, the Feed Grain Program moderated the expanding production of rowcrops in the early 1970s, and the CRP of the 1985 Food Security Act converted cultivated lands into grasslands during the 1980s and 1990s. Ring-necked pheasant populations in Utah (Bartmann 1969) and South Dakota (Erickson and Wiebe 1973) were positively correlated with the amount of cropland retired under the Soil Bank Program. In Oregon, Jarvis and Simpson (1978) reported that peaks in the state's ring-necked pheasant population coincided with years during which the Soil Bank was operative (1956-69). During the Feed Grain Program in Illinois (Joselyn and Warnock 1964) and Wisconsin (Gates and Ostrom 1966) more ring-necked pheasants were produced on program lands, because of higher nesting densities and hatching success, than all other habitat types surveyed. Positive population responses by ring-necked pheasants to cropland converted to grassland for 10-year periods under the CRP were reported in Texas (Berthelsen et al. 1989, Berthelsen et al. 1990), Minnesota (Kimmel et al. 1992), Iowa (Riley 1993), and Ohio (Swanson et al. 1995). In South Dakota, the ring-necked pheasant population dropped from 11 million birds in 1961 during the Soil Bank era to 3 million birds in 1970 after the program was discontinued and grasslands were returned to cultivation (Erickson and Wiebe 1973). When crop retirement programs (Soil Bank and Feed Grain) were phased out in Illinois, nesting ring-necked pheasants concentrated in forage crops, pastures, and strip cover where reproductive success was lower compared to retired cropland because of hay cutting and predation (Warner et al. 1987, Etter et al. 1988). The availability of undisturbed nesting and brood-rearing habitat clearly dictates ring-necked pheasant population levels throughout their range.

Over the past decade, the mean day of first cutting of alfalfa, the most widely planted hay cultivar in the Midwest, has been 3 June, about 10 days earlier than it was in the 1950s (Warner and Etter 1989). The destruction of ring-necked pheasant hens, nests, and chicks by mowing, especially in forage fields, has long been recognized as an important factor limiting production in intensively farmed landscapes (Leedy and Hendershot 1947, Joselyn and Warnock 1964, Warner and Etter 1989).

Ring-necked pheasant nest success in hayfields harvested during the nesting season (mid-Apr to mid-Jul) was 14% compared to 63% in undisturbed fields (Gates and Hale 1975). Lauber (1991) summarized several studies and reported an average ring-necked pheasant nest success rate of 30% in active agricultural lands (hayfields, pastures, small grains, and rowcrops) versus 67% in undisturbed grasslands. Roadside rights-of-way mowed frequently during the growing season were of little value as ring-necked pheasant nesting habitat in Illinois (Warner et al. 1992). Long-term ring-necked pheasant nest success rates averaged 19% on grasslands mowed before 1 August compared to 30% on roadside rights-of-way mowed after 1 August (Warner and Joselyn 1986). Ring-necked pheasant nesting density and nest success were higher in switchgrass (Panicum virgatum) plots that were not mowed or grazed the previous summer than in those that were disturbed (George et al. 1979). Gates and Hale (1975) reported lower ring-necked pheasant nesting densities in fields mowed the year before (52 nests/100 ha) than in fields idled for >/=1 year (101 nests/100 ha). George et al. (1979) and Gates and Hale (1975) attributed the higher ring-necked pheasant nesting densities and production in undisturbed fields to the greater amount of residual cover. In a Wisconsin study, 17% of all successful ring-necked pheasant nests were in grasslands idled >/=1 year that comprised only 4.3% of the study area (Gates and Ostrom 1966).

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