Index of Species Information

WILDLIFE SPECIES:  Poecile atricapillus


WILDLIFE SPECIES: Poecile atricapillus
AUTHORSHIP AND CITATION : Sullivan, Janet. 1995. Poecile atricapillus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: [].
ABBREVIATION : POAT COMMON NAMES : black-capped chickadee TAXONOMY : The currently accepted scientific name for the black-capped chickadee is Poecile atricapillus Linnaeus [2]. The 1957 A.O.U. checklist [1] (the last one that included subspecies) listed nine subspecies of the black-capped chickadee: Poecile atricapillus atricapillus, eastern black-capped chickadee P. a. practicus (Oberholser), Appalachian black-capped chickadee P. a. bartletti Aldrich and Nutt., Newfoundland black-capped chickadee P. a. turneri Ridgway, Yukon black-capped chickadee P. a. septentrionalis Harris, long-tailed chickadee P. a. occidentalis Baird, Oregon chickadee P. a. fortuitus (Dawson and Bowles), Columbian black-capped chickadee P. a. nevadensis (Linsdale), pallid black-capped chickadee P. a. garrinus Behle, Rocky Mountain black-capped chickadee Where the range of the black-capped chickadee overlaps that of other chickadees (Poecile spp.) they are segregated by habitat. There are some areas of breeding territory which the black-capped chickadee and the Carolina chickadee (P. carolinensis) both use; in this area hybrids of the two species commonly occur. There is some disagreement as to the true status of the Carolina chickadee; it has been argued that it is a subspecies of the black-capped chickadee rather than a separate species. The most recent genetic evidence suggests that they are in fact separate species. Hybrids with the mountain chickadee (P. gambeli) have also been reported but are less common than black-capped chickadee-Carolina chickadee hybrids [30]. ORDER : Passeriformes CLASS : Bird FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY


WILDLIFE SPECIES: Poecile atricapillus
GENERAL DISTRIBUTION : The black-capped chickadee is resident from western and central Alaska, most of Canada south of the arctic circle, south to extreme northwestern California, extreme northeastern Nevada, northern New Mexico, central Indiana, and northern New Jersey. At upper elevations in the Appalachians its range extends farther south [6,31]. ECOSYSTEMS : FRES10 White-red-jack pine FRES11 Spruce-fir FRES15 Oak-hickory FRES17 Elm-ash-cottonwood FRES18 Maple-beech-birch FRES19 Aspen-birch FRES20 Douglas-fir FRES21 Ponderosa pine FRES22 Western white pine FRES23 Fir-spruce FRES24 Hemlock-Sitka spruce FRES25 Larch FRES26 Lodgepole pine STATES :

BLM PHYSIOGRAPHIC REGIONS : 1 Northern Pacific Border 2 Cascade Mountains 5 Columbia Plateau 6 Upper Basin and Range 8 Northern Rocky Mountains 9 Middle Rocky Mountains 10 Wyoming Basin 11 Southern Rocky Mountains 12 Colorado Plateau 13 Rocky Mountain Piedmont 14 Great Plains 15 Black Hills Uplift 16 Upper Missouri Basin and Broken Lands KUCHLER PLANT ASSOCIATIONS : K001 Spruce-cedar-hemlock forest K002 Cedar-hemlock-Douglas-fir forest K003 Silver fir-Douglas-fir forest K004 Fir-hemlock forest K005 Mixed conifer forest K008 Lodgepole pine-subalpine forest K010 Ponderosa shrub forest K011 Western ponderosa forest K012 Douglas-fir forest K013 Cedar-hemlock-pine forest K014 Grand fir-Douglas-fir forest K015 Western spruce-fir forest K016 Eastern ponderosa forest K017 Black Hills pine forest K018 Pine-Douglas-fir forest K019 Arizona pine forest K020 Spruce-fir-Douglas-fir forest K021 Southwestern spruce-fir forest K022 Great Basin pine forest K028 Mosaic of K002 and K026 K029 California mixed evergreen forest K081 Oak savanna K082 Mosaic of K074 and K100 K093 Great Lakes spruce-fir forest K094 Conifer bog K095 Great Lakes pine forest K096 Northeastern spruce-fir forest K097 Southeastern spruce-fir forest K098 Northern floodplain forest K099 Maple-basswood forest K100 Oak-hickory forest K101 Elm-ash forest K109 Transition between K104 and K106 SAF COVER TYPES : 1 Jack pine 5 Balsam fir 12 Black spruce 13 Black spruce-tamarack 14 Northern pin oak 15 Red pine 16 Aspen 17 Pin cherry 18 Paper birch 19 Gray birch-red maple 20 White pine-northern red oak-red maple 21 Eastern white pine 22 White pine-hemlock 23 Eastern hemlock 24 Hemlock-yellow birch 25 Sugar maple-beech-yellow birch 26 Sugar maple-basswood 27 Sugar maple 28 Black cherry-maple 30 Red spruce-yellow birch 31 Red spruce-sugar maple-beech 32 Red spruce 33 Red spruce-balsam fir 34 Red spruce-Fraser fir 35 Paper birch-red spruce-balsam fir 37 Northern white-cedar 38 Tamarack 39 Black ash-American elm-red maple 40 Post oak-blackjack oak 42 Bur oak 43 Bear oak 44 Chestnut oak 46 Eastern redcedar 50 Black locust 51 White pine-chestnut oak 52 White oak-black oak-northern red oak 53 White oak 55 Northern red oak 57 Yellow-poplar 58 Yellow-poplar-eastern hemlock 59 Yellow-poplar-white oak-northern red oak 60 Beech-sugar maple 62 Silver maple-American elm 63 Cottonwood 87 Sweetgum-yellow-poplar 93 Sugarberry-American elm-green ash 94 Sycamore-sweetgum-American elm 95 Black willow 107 White spruce 108 Red maple 109 Hawthorn 110 Black oak 201 White spruce 202 White spruce-paper birch 204 Black spruce 205 Mountain hemlock 206 Engelmann spruce-subalpine fir 210 Interior Douglas-fir 208 Whitebark pine 210 Interior Douglas-fir 211 White fir 212 Western larch 213 Grand fir 215 Western white pine 216 Blue spruce 218 Lodgepole pine 219 Limber pine 223 Sitka spruce 224 Western hemlock 225 Western hemlock-Sitka spruce 227 Western redcedar-western hemlock 228 Western redcedar 229 Pacific Douglas-fir 230 Douglas-fir-western hemlock 234 Douglas-fir-tanoak-Pacific madrone 237 Interior ponderosa pine 238 Western juniper 244 Pacific ponderosa pine-Douglas-fir 245 Pacific ponderosa pine 251 White spruce-aspen 253 Black spruce-white spruce 254 Black spruce-paper birch SRM (RANGELAND) COVER TYPES : 109 Ponderosa pine shrubland 110 Ponderosa pine-grassland 412 Juniper-pinyon woodland 422 Riparian PLANT COMMUNITIES : Black-capped chickadee habitat includes evergreen forested wetlands, deciduous forested wetlands [35], deciduous woodlands, mixed woodlands, deciduous and coniferous forests, orchards, deciduous shrubs, urban and suburban areas [6], and disturbed areas such as old fields. Favored riparian communities include cottonwood (Populus spp.) and sometimes willow (Salix spp.) thickets. Birches (Betula spp.) and alders (Alnus spp.) are often used for both food and nesting, but black-capped chickadees use a wide variety of other plant species as well. Black-capped chickadees occur in many habitat types [31]. In western North Dakota black-capped chickadees forage in the canopy and nest in cavities in cottonwood stands; they occupy ash (mostly green ash [Fraxinus pennsylvanica]) woodland interiors, and are found in pine (Pinus spp.) communities. Black-capped chickadees are common in dense ponderosa pine (P. ponderosa) stands with well-developed shrub layers [14]. In the Konza Prairie, Kansas, black-capped chickadees were the second most abundant species in gallery forests. These forests are dominated by bur oak (Quercus macrocarpa), chinkapin oak (Q. muehlenbergii), and hackberry (Celtis occidentalis) [8]. Black-capped chickadees also occur on adjacent prairie dominated by big bluestem (Andropogon gerardii), little bluestem (Schizachyrium scoparium), switchgrass (Panicum virgatum), and indiangrass (Sorghastrum nutans). The presence of black-capped chickadees in grassland habitat was attributed to the availability of isolated tree and shrub patches along ravines [8]. In Illinois black-capped chickadees were recorded in northern red oak (Q. rubra)-sugar maple (Acer saccharum)-hackberry dominated woodlands [35].


WILDLIFE SPECIES: Poecile atricapillus
TIMING OF MAJOR LIFE HISTORY EVENTS : Pair Formation: The peak period for pair formation is fall and is associated with winter flock formation; winter flocks consist largely of mated pairs. Even very young black-capped chickadees form pair bonds at this time. A few pairs are formed in winter following mortality of high-ranking members of the flock, and occasionally floaters (low-ranking individuals unattached to a mate or a flock) establish new pair bonds in spring [30,31]. Territory Establishment: Territories are established in spring, during winter flock break-up; this period varies with area, year, and other factors. Breeding territory boundaries are usually established 5 to 7 weeks before onset of egg-laying [30,31]. Nesting: Both the male and female excavate the nest hole, but the female builds the nest. The cup-shaped nest consists of cottony plant fibers, hairs, wool, moss, and leaves, and is lined with hair, plant down, wool, and feathers [12]. Clutch: Eggs are laid from early April to mid-July depending on spring weather and food availability [30]. Usually 1 egg is laid per day; the average clutch size is from 6 to 8 eggs, ranging from 5 to 10. Eggs are incubated for 12 to 13 days by the female, who is fed by the male [38]. All eggs usually hatch within 12 to 30 hours of each other, usually in the order laid [31]; nestlings are present from early May to late July [30,31]. Development of Young: Black-capped chickadees are altricial; newly hatched young are blind and nearly naked. They have pinfeathers by about day 9, and usually fledge on day 16. When nests are disturbed, fledglings may leave the nest early, sometimes as early as 12 days. Fledglings are fed by the parents for 2 to 4 weeks (3-4 weeks is typical) [30,31]. Longevity: Age at first breeding is typically less than 1 year, although some individuals may not breed until they are 1 or 2 years old. The average lifespan of black-capped chickadees is approximately 2.5 years; however, 5-year-old birds were not uncommon in northwestern Connecticut [17]. The longest lived black-capped chickadee on record was at least 12 years 5 months at the time of last banding [31]. Mortality: Black-capped chickadees are fairly cold hardy; the majority of black-capped chickadee mortality is believed to be caused by winter malnutrition, which reduces the ability to withstand cold weather and resist disease. There have been few major outbreaks of diseases in black-capped chickadee populations and there are relatively few nest parasites [31]. Wintering: Wintering flocks of black-capped chickadees usually consist of four to eight individuals [9]. Black-capped chickadee residence in cold climates is made possible by night torpor, a regulated hypothermia which allows black-capped chickadees to survive cold nights with minimum energy loss [31]. Seasonal Movements: Long-distance movements are usually only undertaken by black-capped chickadees less than 1 year old. However, large numbers of black-capped chickadees emigrate at irregular intervals of about 2 years. These movements are more properly termed irruptions than seasonal migrations. Factors influencing irruptions, particularly in the eastern portions of the black-capped chickadee range, include fluctuation in northern seed crops and unusually high recruitment rates. Fall movements tend to be south or southwest; spring movements are usually northward but are sometimes aimless [31]. Movement over water is avoided or undertaken only with great hesitation [30,38]. PREFERRED HABITAT : Black-capped chickadees prefer relatively open sites near deep woods [6]. They are usually more common near edges, but also occur in the interior of wooded tracts [31]. In Iowa most observations of black-capped chickadees were in floodplain woodlands and scrub; fewer black-capped chickadees were observed in upland woodlands, wooded edges, and savannah (in descending order of numbers of observations). Black-capped chickadee observations were positively correlated with sapling and tree species richness, sapling and tree size, and vertical patchiness. There was a negative correlation with vine density and with snag hardness [32]. In Saskatchewan black-capped chickadees were found in aspen (Populus spp.) groves larger than 0.5 acre (0.2 ha) in area, and did not occur in smaller groves [15]. In Montana foliage-insect feeders including black-capped chickadees were observed most often in uncut forests. Black-capped chickadees fed primarily where foliage canopy was well developed above 26.4 feet (8 m) [24]. Riparian communities are important to black-capped chickadees and other gleaners (birds which search vegetation for stationary prey). Emerging aquatic insects are a particularly valuable food for gleaners. Mayflies and stoneflies spend most of the daylight hours resting on low vegetation near the stream channel. The density of gleaners (in this area black-capped chickadees were the most abundant gleaners) is positively correlated with emergence rates of aquatic insects [10]. In Colorado black-capped chickadee nest site selection was positively associated with density of small trees; in Missouri and Tennessee, black-capped chickadees are reported to prefer small trees and young open forest [28]. COVER REQUIREMENTS : Nesting: Black-capped chickadees excavate holes in soft decayed wood and also use existing cavities (but usually only if there is material to be excavated) [30,31]. Dead standing trees greater than 4 inches (10 cm) dbh are used for nesting and feeding [6]. In Illinois nests were found in cavities of stubs (broken off snags). The stubs were usually 5 to 6.6 feet (1.6-2 m) tall and 4.3 to 5.1 inches (11-13 cm) in diameter [41]. Common nest sites are stubs of gray birch (Betula populifolia) or paper birch (B. papyrifera) [12], but almost any early seral species with soft wood may be used; the particular tree species favored depends on the region. Most of these trees occur as living trees in early seral stages, are short-lived, and persist into intermediate seral stages as decaying snags [40]. For example, Odum [23] reported that of 18 black-capped chickadee nests he observed in upstate New York, 4 were in pin cherry (Prunus pensylvanica), 3 in paper birch, 3 in American beech (Fagus grandifolia), 2 in yellow birch (Betula alleghaniensis), 2 in willows (Salix spp.), 1 in basswood (Tilia americana), 1 in sugar maple, 1 in white ash (Fraxinus americana), and 1 in an apple tree (Malus spp.) [40]. In Vermont northern hardwoods forests, most black-capped chickadee nest trees were in an advanced state of decay with soft outer wood. Most nests were in trees that were shorter than neighboring non-nest trees, but no smaller in diameter [26]. Nest trees used by black-capped chickadees in northwestern Montana western larch (Larix occidentalis)-Douglas-fir (Pseudotsuga menziesii) forests averaged 8 inches (20 cm) dbh and ranged from 4 to 12 inches (10-30.5 cm) dbh. This was the smallest average diameter used by any of the cavity-nesting birds observed [21]. Most nests were in broken-topped larch trees [20]. In Iowa 92 percent of black-capped chickadee nests in riparian communities were in snags, 4 percent were in dead limbs of living trees, and 4 percent were in living trees. There was a positive association between black-capped chickadee use and snag size in snags less than 9 inches (25 cm) dbh [32]. In a riparian area in Colorado with a viable black-capped chickadee population, snags are not plentiful but large dead branches are. In an area dominated by plains cottonwood (Populus deltoides var. occidentalis) with some peachleaf willow (S. amygdaloides) and boxelder (Acer negundo), cottonwood snags comprised 2.7 percent of all cottonwood stems. This density of snags is quite low, primarily due to the decadence of the stand. However, limb trees (trees with more than 3.3 feet [1 m] of dead limbs greater than 4 inches (10 cm) in diameter) made up 47 percent of the cottonwood population [27]. In western Montana, McClelland and others [19] observed black-capped chickadees using cavities excavated by sapsuckers (Sphyrapicus spp.) [19]. Birdhouses are used occasionally [6]. Roosting: Black-capped chickadees roost primarily in thick vegetation or in cavities, particularly on cold nights. Flocks seldom roost clumped together, but flock members usually roost near each other [31]. Foraging and Feeding: Flock members usually feed from 3.3 to 33 feet (1-10 m) apart, occasionally feeding within 2.5 inches (6 cm) for brief periods [31]. Breeding Territory: Black-capped chickadee breeding territory size varies with habitat quality, black-capped chickadee population density, rank, and the course of the breeding season [30]. In upstate New York breeding territories ranged in size from 8.4 acres (3.4 ha) to 17.1 acres (6.9 ha) and averaged 13.2 acres (5.3 ha) [23]. An eastern Massachusetts population had an average breeding territory size of 10.7 acres (4.3 ha), but ranged from 3.8 to 17.9 acres (1.5-7.2 ha) [30]. FOOD HABITS : Foraging: Black-capped chickadees forage from ground to treetop; ground foraging birds have usually been displaced by higher ranking birds [30]. Black-capped chickadees forage on tree trunks, branches, and foliage [11], feeding on insects [34], seeds, and berries. Five basic foraging maneuvers used by black-capped chickadees are 1) gleaning (57% of time spent foraging), 2) hanging from leaf or twig to capture food items (28%), 3) hovering (8.8%), 4) probing (3.5%), and 5) catching insects in flight, called hawking (2.4%). These proportions probably vary with availability of prey, season, and other factors [31]. Caching: Black-capped chickadees cache seeds from open cones [11]. Insects are also cached. Most caching occurs in the fall, but caching may occur at any time food is plentiful. Storage sites include bark, dead leaves, clusters of conifer needles, dirt, and snow. Black-capped chickadees scatter-hoard; they hide each individual food item in a separate spot [31]. Sherry [29] reported that black-capped chickadees can remember cache sites for at least 24 hours, and Hitchcock and Sherry [13] reported that captive black-capped chickadees can recover caches after 28 days. Animal Foods: In winter, approximately 50 percent of black-capped chickadee foods are animal foods, the rest seeds and berries. During the breeding season, 80 to 90 percent of the black-capped chickadee diet is animal foods [18,31]. Winter black-capped chickadee animal foods consist mostly of eggs of moths, plant lice, katydids, and spiders. In summer moths, caterpillars, spiders, beetles (particularly weevils), flies, wasps, true bugs, plant lice, scale insects, leafhoppers, and tree hoppers are common food items [18]. Smith [30] described the black-capped chickadee summer diet as consisting largely of caterpillars, including some hairy caterpillars such as early instar gypsy moths and tent caterpillars. Black-capped chickadees have been observed taking animal fat from carrion and eating suet and peanut butter at feeders [30]. Plant Foods: Black-capped chickadee plant foods are mainly seeds and berries including goldenrods (Solidago spp.) and staghorn sumac (Rhus typhina) seeds in fall. Pine seeds are a main staple in fall, winter, and spring. Seeds of hemlocks (Tsuga spp.) and birches are eaten in winter; seeds or fruits of poison-ivies (Toxicodendron spp.), blueberries and huckleberries (Vaccinium spp.), bayberries (Myrica spp.), ragweeds (Ambrosia spp.), sunflowers (Helianthus spp.), chokecherry (Prunus virginiana), and Virginia creeper (Parthenocissus quinquefolia) are eaten in spring and summer [18]. Raspberries (Rubus spp.), cherries (Prunus spp.), and yellow-poplar (Liriodendron tulipifera) seeds are also consumed [30]. Fluids: Black-capped chickadees drink when water is available. Fluids are derived mostly from foods in winter; highest demand for liquid water is in summer [31]. PREDATORS : Natural predators of the black-capped chickadee include goshawk (Accipiter gentilis), sharp-shinned hawk (A. striatus), Cooper's hawk (A. cooperii), American kestrel (Falco sparverius), merlin (F. columbarius), northern pygmy-owl (Glaucidium gnoma), and northern shrike (Lanius excubiter). Around birdfeeders, the black-capped chickadee is often preyed on by the domestic cat (Felis catus). Nest predators are largely excluded by the small size of black-capped chickadee nest entrance holes, but very small squirrels (Tamiascurius spp.) or chipmunks (Tamias spp.) occasionally raid black-capped chickadee nests. Weasels (Mustela spp.) and climbing snakes pose a threat to eggs and nestlings [30]. MANAGEMENT CONSIDERATIONS : A review of Christmas Bird Count data for the Pacific Northwest showed that most black-capped chickadee populations have apparently been stable during the past 40 years. Of 49 locales reporting, 6 showed significant declines (stations in Alaska, British Columbia, Montana, and Oregon), 5 showed significant increases (British Columbia, California, Montana, and Washington), and the remaining 38 showed no overall change. It was speculated that the "significant" increases and decreases may actually represent anomalous data [4]. Long-term wildlife management should strive for sites with a mosaic of age structures [28]. In northwestern Connecticut the clearcutting of 60 acres (24.4 ha) of red pine (Pinus resinosa) within a 321 acre (130 ha) banding plot had no discernible effect on black-capped chickadee populations in a long-term population study [17]. Forest clearing can increase edge, which is preferred (but not required) black-capped chickadee habitat. Removal of snags and cull trees with dead limbs decreases available nest sites for black-capped chickadees [31,32], although black-capped chickadees are listed as tolerant of habitat alteration [32]. Stauffer and Best [32] listed the following predicted effects of various types of habitat alteration on black-capped chickadee populations: removal of all wood vegetation: elimination reduce woody vegetation to narrow strips: negative woody canopy partly removed: no effect woody canopy partly removed, shrubs and saplings thinned: negative shrubs and saplings thinned: negative snags removed: negative Lack of cottonwood regeneration is detrimental to the long-term stability of cavity-nesting bird populations [17]. In cold-winter areas, feeders often enhance black-capped chickadee survival, particularly in disturbed areas where food supplies are limited [31]. Nest boxes can increase available nest sites where natural cavities are limited. Nest boxes are not readily used unless they are half-filled with sawdust, apparently so that the birds have something to excavate [30]. Black-capped chickadees are only rarely host to the brown-headed cowbird (Molothrus ater) [31]. Black-capped chickadees are important predators of larch casebearer larvae and pine sawfly larvae [5].


WILDLIFE SPECIES: Poecile atricapillus
DIRECT FIRE EFFECTS ON ANIMALS : There is no information in the published literature concerning direct mortality of black-capped chickadees from fire. It is likely that, like most birds, black-capped chickadees can easily escape fire; any substantial mortality is likely to occur only when severe fires occur during the early breeding season. HABITAT RELATED FIRE EFFECTS : The most likely change due to fire in black-capped chickadee habitat is with respect to snag density (nest site availability) and food availability. Niemi [22] suggested that chickadee populations generally decrease following fire, probably due to a decrease in habitat complexity and available food. In northern Rocky Mountain conifer forests that were severely burned within 1 to 2 years of the study, black-capped chickadees were detected on 13 of 33 sites. In a survey of bird habitat studies in northern Rocky Mountain dryland habitats, segregated by habitat type, black-capped chickadees were found on early successional (less than 10 years old) burned forest (48% of 23 studies), and mid-successional (10 to 40 years old) burned forest (40% of 5 studies). Studies reporting observations of birds in cottonwood bottomlands had the highest proportion of black-capped chickadee observations (64% of 21 studies). Black-capped chickadees had a habitat preference average of 9.71 out of 15 possible habitats (if this figure were 1, a bird species is restricted to only 1 of the 15 habitats, if the figure were 15, the species has shown absolutely no preference for any of the available habitat types) [39]. In northern Minnesota the Little Sioux fire burned 15,000 acres (6,072 ha) of northern hardwoods and pine forests. Black-capped chickadees were common on unburned stands, but in postfire years 2, 3, and 4 they were uncommon on all burned study sites [22]. In north-central Colorado a severe 1966 wildfire in lodgepole pine (Pinus contorta) with subalpine fir (Abies lasiocarpa) and Engelmann spruce (Picea engelmannii) resulted in widespread crown mortality. In 1974 there were many standing dead trees on the burned site. There were no black-capped chickadees observed on the burned site, but there were a few in the ecotone and in adjacent unburned lodgepole stands [25]. FIRE USE : NO-ENTRY


WILDLIFE SPECIES: Poecile atricapillus
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[26015] 13. Hitchcock, Christine L.; Sherry, David F. 1990. Long-term memory for cache sites in the black-capped chickadee. Animal Behavior. 40: 701-712. [25114] 14. Hopkins, Rick B.; Cassel, J. Frank; Bjugstad, Ardell J. 1986. Relationships between breeding birds and vegetation in four woodland types of the Little Missouri National Grasslands. Res. Pap. RM-270. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 12 p. [2758] 15. Johns, Brian W. 1993. The influence of grove size on bird species richness in aspen parklands. Wilson Bulletin. 105(2): 256-264. [22269] 16. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society. 77 p. [1384] 17. Loery, Gordon; Nichols, James D. 1985. Dynamics of a black-capped chickadee population, 1958-1983. Ecology. 66(4): 1195-1203. [24338] 18. 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