Melanerpes erythrocephalus



INTRODUCTORY


  Photo by: Stephen Cresswell

AUTHORSHIP AND CITATION:
Luensmann, Peggy. 2006. Melanerpes erythrocephalus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: Available: http://www.fs.fed.us/database/feis/ [].

FEIS ABBREVIATION:
MEER

SYNONYMS:
None

COMMON NAMES:
red-headed woodpecker

TAXONOMY:
The scientific name of the red-headed woodpecker is Melanerpes erythrocephalus L. It is a member of the woodpecker family, Picidae [1,79]. Subspecies include [1]:

Melanerpes erythrocephalus erythrocephalus L.
Melanerpes erythrocephalus caurinus Brodkorb

ORDER:
Piciformes

CLASS:
Bird

FEDERAL LEGAL STATUS:
None

OTHER STATUS:
Information on state-level protected status of animals in the United States is available at NatureServe, although recent changes in status may not be included.

ANIMAL DISTRIBUTION AND OCCURRENCE

SPECIES: Melanerpes erythrocephalus
GENERAL DISTRIBUTION:
The range of the red-headed woodpecker extends from extreme southern Quebec and Ontario south to Florida and west to the Rocky Mountains. Occasional vagrants may travel outside this range [6,65].

ECOSYSTEMS [32]:
FRES10 White-red-jack pine
FRES11 Spruce-fir
FRES12 Longleaf-slash pine
FRES13 Loblolly-shortleaf pine
FRES14 Oak-pine
FRES15 Oak-hickory
FRES16 Oak-gum-cypress
FRES17 Elm-ash-cottonwood
FRES18 Maple-beech-birch
FRES30 Desert shrub
FRES32 Texas savanna
FRES33 Southwestern shrubsteppe
FRES35 Pinyon-juniper
FRES38 Plains grasslands
FRES39 Prairie
FRES40 Desert grasslands

STATES/PROVINCES: (key to state/province abbreviations)
UNITED STATES
AL AR CO DE FL GA IL IN IA KS
KY LA MD MA MI MN MS MO MT NE
NJ NM NY NC ND OH OK PA SC SD
TN TX VA WV WI WY DC

CANADA
MB ON

BLM PHYSIOGRAPHIC REGIONS [8]:
13 Rocky Mountain Piedmont
14 Great Plains
16 Upper Missouri Basin and Broken Lands

KUCHLER [52] PLANT ASSOCIATIONS:
K081 Oak savanna
K082 Mosaic of K074 and K100
K083 Cedar glades
K084 Cross Timbers
K089 Black Belt
K090 Live oak-sea oats
K098 Northern floodplain forest
K099 Maple-basswood forest
K100 Oak-hickory forest
K101 Elm-ash forest
K102 Beech-maple forest
K103 Mixed mesophytic forest
K104 Appalachian oak forest
K106 Northern hardwoods
K107 Northern hardwoods-fir forest
K108 Northern hardwoods-spruce forest
K109 Transition between K104 and K106
K110 Northeastern oak-pine forest
K111 Oak-hickory-pine
K112 Southern mixed forest
K113 Southern floodplain forest
K114 Pocosin
K115 Sand pine scrub
K116 Subtropical pine forest

SAF COVER TYPES [30]:
14 Northern pin oak
20 White pine-northern red oak-red maple
25 Sugar maple-beech-yellow birch
26 Sugar maple-basswood
27 Sugar maple
28 Black cherry-maple
31 Red spruce-sugar maple-beech
39 Black ash-American elm-red maple
40 Post oak-blackjack oak
42 Bur oak
43 Bear oak
44 Chestnut oak
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
65 Pin oak-sweetgum
69 Sand pine
70 Longleaf pine
71 Longleaf pine-scrub oak
72 Southern scrub oak
73 Southern redcedar
74 Cabbage palmetto
75 Shortleaf pine
76 Shortleaf pine-oak
78 Virginia pine-oak
80 Loblolly pine-shortleaf pine
81 Loblolly pine
82 Loblolly pine-hardwood
83 Longleaf pine-slash pine
84 Slash pine
85 Slash pine-hardwood
87 Sweetgum-yellow-poplar
88 Willow oak-water oak-diamondleaf (laurel) oak
89 Live oak
91 Swamp chestnut oak-cherrybark oak
92 Sweetgum-willow oak
93 Sugarberry-American elm-green ash
94 Sycamore-sweetgum-American elm
95 Black willow
96 Overcup oak-water hickory
98 Pond pine
108 Red maple
110 Black oak
111 South Florida slash pine
235 Cottonwood-willow
236 Bur oak

SRM (RANGELAND) COVER TYPES [78]:
731 Cross timbers-Oklahoma
732 Cross timbers-Texas (little bluestem-post oak)
801 Savanna
802 Missouri prairie
803 Missouri glades
804 Tall fescue
805 Riparian
808 Sand pine scrub
809 Mixed hardwood and pine
810 Longleaf pine-turkey oak hills
811 South Florida flatwoods
812 North Florida flatwoods
814 Cabbage palm flatwoods
815 Upland hardwood hammocks
816 Cabbage palm hammocks
817 Oak hammocks

PLANT COMMUNITIES:
See lists above.

BIOLOGICAL DATA AND HABITAT REQUIREMENTS

SPECIES: Melanerpes erythrocephalus
TIMING OF MAJOR LIFE HISTORY EVENTS:
Red-headed woodpeckers are temperate migrants with migratory habits that are largely influenced by the availability of mast in winter [6,40,63]. Migrants typically travel south and east in autumn if the mast fails at breeding sites. If mast is plentiful, however, red-headed woodpeckers often overwinter at or near their breeding sites [80]. Fall migration occurs from late August to early November with a peak in September [6,35,63,80]. Spring migration commences in mid-February with a peak from late April to mid-May [6,35,80].

Red-headed woodpeckers are highly territorial and aggressive [6,49,63]. Confrontations between conspecifics and other species are common [6,42,69]. Red-headed woodpeckers usurp nests from northern flickers (Colaptes auratus) and red-bellied woodpeckers (Melanerpes carolinus) [42,45]. In turn, red-headed woodpecker nests are occasionally usurped by European starlings (Sturnus vulgaris) [42,43,45]. Despite their aggressive temperaments, red-headed woodpeckers occasionally share a nest tree with other species including red-bellied woodpeckers, northern flickers, European starlings, American kestrels (Falco sparverius), and Indiana bats (Myotis sodalis) [43,44,53,69,94].

Although solitary in winter [57,62], red-headed woodpeckers are monogamous and may remain paired for several breeding seasons [44]. Red-headed woodpeckers are primary cavity nesters that excavate their own nest and roost cavities [80,83]. However, they also use existing holes for nesting and roosting [5,20,80]. Nesting is typically initiated from early May to mid-June, but it may begin as early as February in the Southeast [42,45,69,85]. Excavation of nest cavities begins during the second half of April, peaks in early May, and continues until the end of July [35,42]. Egg laying begins in early May and usually ends by mid-August [7,35,42,48]. Clutches contain 4 to 5 eggs on average, with 3 to 10 eggs possible [6,35,42,48,59,96]. Some pairs raise 2 broods in a season [6,42,43,44]. Clutch size is typically larger during first brood attempts than second brood attempts [42].

Incubation lasts 12 to 14 days [6,7,48,59,69]. Asynchronous hatching may occur if incubation begins before all eggs are laid [48]. Nestlings are present from mid-May to late August with a peak in early to mid-June in the southern extent of the range and a mid-July peak in the northern extent [42,45]. Fledging occurs from the second week of June to the first week of September [7,35,42,45]. Nestlings fledge at approximately 24 to 30 days of age [6,7,59,69]. Pairs in Mississippi fledged 2.1 to 2.3 young/brood on average [42]. Parents drive juveniles from the nest several weeks after fledging [7,62].

Red-headed woodpeckers generally have high nesting success. In Colorado, red-headed woodpeckers experienced 50% to100% nesting success [7]. Average nest success was 78% throughout the United States [59].

Red-headed woodpecker mortality is highly variable. Winter mortality of red-headed woodpeckers in Ohio was 7% [23]. Annual mortality of adult red-headed woodpeckers was reported as 38% in a review [59]. Adult mortality was low in a Colorado study [7]. Red-headed woodpeckers may succumb to exposure in winter. Both adult and juvenile red-headed woodpeckers were found dead in Illinois after heavy snowfall and severe cold. Immature red-headed woodpeckers may fare worse than adults [34].

The maximum life span of the red-headed woodpecker is unknown. One red-headed woodpecker was recaptured 9 years and 11 months after banding [12]. However, most red-headed woodpeckers probably do not live more than 2 years [35].

PREFERRED HABITAT:
Primary habitats used by red-headed woodpeckers include oak (Quercus spp.) savanna and mature open bottomland forest [10,15,35,63,69,83]. Red-headed woodpeckers are also found in upland forests, woodlots, shelterbelts along agricultural fields, residential areas, golf courses, and other habitats containing mature open hardwoods with snags or trees with dead limbs [14,35,38,58,63,69,71,83]. The use of herbaceous habitats or stands with high canopy cover or a dense mid-story is infrequent [38,83]. During periods of cold and deep snow cover, red-headed woodpeckers may move from the bottomlands to the uplands [34].

Red-headed woodpeckers show a preference for mature open canopy forests with large trees, a high basal area, and little understory [14,15]. In a Texas study, 79% of red-headed woodpeckers were observed in a bottomland hardwood forest that had high crown closure in the overstory and low understory and mid-story growth. Hardwood-dominated ridges adjacent to the bottomland forest housed nearly 17% of the red-headed woodpeckers observed. Mixed pine (Pinus spp.)-hardwood forests and pine uplands were occupied by approximately 2% of the observed red-headed woodpeckers each [18]. Red-headed woodpeckers frequent savannas and forest edges during the breeding season, and mostly avoid the interior of forest patches >30 acres unless fire, wind damage, or some other event creates gaps in the forest interior [35]. In Illinois, red-headed woodpeckers preferred the forest edge from late summer to early winter and the interior forest during the rest of the year [91]. Red-headed woodpecker nests in Ohio were associated with low ground cover within a mosaic of agricultural and forested habitats [46]. Oak woodlots with an abundance of "overmature" trees with dead branches and cavities support red-headed woodpeckers during years of high acorn mast [63]. In South Dakota, red-headed woodpeckers were observed in riparian habitat adjacent to black-tailed prairie dog (Cynomys ludovicianus) colonies [77].

Mean habitat characteristics of a plains cottonwood-peachleaf willow (Populus deltoides ssp. monilifera-Salix amygdaloides) riparian stand in Colorado and 2 eastern cottonwood/Rocky Mountain juniper (P. deltoides/Juniperus scopulorum) stands in North Dakota inhabited by red-headed woodpeckers were:

Location Basal area
(m/ha)
Canopy cover
(%)
Mean canopy height
(m)
Large tree density
(trees/ha)
Medium tree density
(trees/ha)
Small tree density
(trees/ha)
North Dakota 15.3-18.5 70.0-85.0 20.0-21.0 7.7-14.8 95.8-103.7 79.0-82.0 [41]
Colorado 11.3 26.6 .... 8.1 75.0 54.8 [75]

Red-headed woodpecker habitat in a bottomland hardwood forest in Texas had an average of 4.3 trees greater than 15 inches (38 cm) DBH/0.04 ha and a dominant vegetation height of 108.9 feet (33.2 m) [16]. In Illinois, there was a positive relationship between the number of red-headed woodpeckers and the number of oaks greater than 10 inches (25 cm) DBH (r=0.909, P<0.05 in year 1; r=0.961, P<0.01 in year 2) [35]. More than half of all plains cottonwoods in a Colorado stand had an average DBH of 9 to 20 inches (24-52 cm) [74]. Optimal stand structure for red-headed woodpeckers in longleaf pine-loblolly pine (Pinus palustris-P. taeda) stands in South Carolina would likely be DBH of 16 to 24 inches (40-61 cm) for cavity trees [39]. Red-headed woodpeckers in Iowa responded negatively (R=0.113, P≤0.01) to tree density but positively (R=0.113, P≤0.01) to snag density. Red-headed woodpeckers also responded positively (R=0.113, P≤0.01) to tree size and forb species richness. Snags with an average DBH of 20 to 30 inches (51-75 cm) were highly favored (R=0.113, P≤0.01) [83].

In late seral stage stands in South Dakota, little mid-story vegetation was present. Average vegetation characteristics of late seral and late-intermediate seral stands in the plains cottonwood/willow (Salix spp.) riparian habitats occupied by red-headed woodpeckers were [72]:

  Late intermediate seral stage Late seral stage
% cover of vegetation <1 m tall 54.6 79.9
Number of cottonwood seedlings/acre 123.5 2.5
Number of willow seedlings/ha 46.2 8.4
Number of cottonwood trees/ha 949.7 116.5
Number of small snags/ha 52.1 0.0
Number of large snags/ha 0.7 6.1
Basal area of cottonwoods (m/ha) 332.6 766.0
DBH of cottonwoods (cm) 17.0 67.3
% overstory canopy cover 58.7 48.5

Snags are essential habitat components that are utilized for nesting, roosting, and foraging [35,38,69]. American beaver (Castor canadensis) ponds, open wooded swamps, rivers bordered by hardwoods, and other riparian or wetland areas where snags are plentiful provide valuable habitat [69,97]. Similarly, trees killed by flooding as well as those bordering lakes, marshes, and other frequently flooded areas support red-headed woodpecker populations [63]. In addition, red-headed woodpeckers are highly attracted to forested areas that have been treated with herbicides and contain numerous standing snags [38]. Snag densities averaged 0.66/ha in a plains cottonwood-dominated stand in Colorado [74]. An average of 2.7 snags was present within 0.04-ha plots in a Texas bottomland hardwood forest. Average snag DBH was 9.5 inches (24.1 cm) [16].

Red-headed woodpeckers utilize live and dead trees for foraging [16]. Foraging activities most often occurred in association with live trees (48.5%) followed by nest snags (29.5%) and other snags (22.1%) in Florida [94]. Mean foraging height for the red-headed woodpecker depends on the condition of the trees used. In a Texas study, mean foraging height in live trees was 43 feet (13 m); in dead snags red-headed woodpeckers foraged at 39 feet (12 m) on average; and mean forage height was 41 feet (12.5 m) in live trees with dead limbs [16]. Red-headed woodpecker activity was strongly restricted to the canopy, between 24.9 feet (7.6 m) above ground level to the canopy top, in a Louisiana bottomland hardwood forest [22].

Territory size/density: Small forest tracts are utilized if the habitat provides adequate nesting sites, roosting sites, and food. Red-headed woodpeckers in a South Carolina bottomland hardwood forest were most commonly detected in 0.5-ha gaps and least common in 0.06-ha gaps. Differences were not significant (P=0.26) [60]. In Arkansas and Oklahoma, no red-headed woodpeckers were observed in forest gaps ranging 0.54 to 2.62 acres in size. However, red-headed woodpeckers were detected at an average density of 0.3 bird/acre in clearcuts 35 to 40 acres in size [88]. In Virginia, red-headed woodpeckers nested in oak woodlots as small as 2 ha, while oak woodlots as small as 3 ha supported 2 breeding pairs [14].

Red-headed woodpeckers were observed at a rate of 0.59 bird/ha in late-seral cottonwood forests in South Dakota and at 0.05 bird/ha in late intermediate-seral forests during the breeding season [72]. The difference was significant (P≤0.15) [72]. No red-headed woodpeckers were detected in the early and early intermediate-seral stages in South Dakota [72].

Red-headed woodpecker densities and territory sizes vary greatly depending on location, habitat, and season. Densities during the breeding season are reported in the following table:

Location Habitat type Average number of individuals Average number of breeding pairs Area
Colorado mature plains cottonwood bottomland forest 9-16 .... 100 ha [74,76]
plains cottonwood floodplain 20.0 .... 40 ha [7]
Illinois urban and suburban residential 0-25 .... 100 acres
oak-maple (Acer spp.) forest 3-6.0 .... 100 acres
grazed bottomland woods 4 .... 100 acres
mature bottomland forest 3.4 .... 100 acres
mature upland forest 0.4 .... 100 acres
second-growth hardwood forest 5.9-27 .... 100 acres
savanna 18 .... 100 acres
orchards 0-4.4 .... 100 acres
shrublands 0-28 .... 100 acres
swamp prairie 14.3 .... 100 acres
pasture 0-5.9 .... 100 acres
oak-maple forest edge 13.7 .... 1 mile of edge [35]
Iowa tallgrass prairie, mowed edge 1.15 .... 1,000 m of edge
tallgrass prairie, burned edge 0.80 .... 1,000 m of edge
tallgrass prairie, untreated edge 0.36 .... 1,000 m of edge [93]
Louisiana water oak (Q. nigra)-loblolly pine-spruce pine (P. glabra) 11.0 .... 100 acres [61]
North Dakota eastern cottonwood/Rocky Mountain juniper floodplain .... 4.3 40 ha [41]
Oklahoma shortleaf pine (P. echinata)-mixed oak upland forest .... 6.6 100 acres [11]
South Carolina longleaf pine-loblolly pine .... 2.3 100 acres [39]

Breeding territories in Florida ranged from 3.1 to 8.5 ha [94]. Average territory size in Illinois was 5.5 to 11.0 ha [91].

Red-headed woodpecker densities during winter are reported in the following table:

Location

Habitat type

Average number of individuals Area
Illinois urban residential 0-0.5 100 acres
oak-maple forest 12.5 100 acres [35]
mature bottomland forest 33.7-34.2 100 acres
mature upland forest 8.2-8.4 100 acres [34,35]
grazed bottomland forest 1.3 100 acres
shrubby field 0-7 100 acres
oak-maple forest edge 6.8 1 mile of edge [35]
Maryland pin oak-ash-black locust (Q. palustris-Fraxinus spp.-Robinia pseudoacacia) 12 1.25 ha [49]

Actual red-headed woodpecker population densities across all winter habitats in Illinois ranged from 0 to 89 individuals/100 acres [35].

Winter territories in an Ohio beech (Fagus spp.) grove averaged 0.05 ha for adults and 0.03 ha for juveniles [23]. In Florida, winter territories averaged 1.00 ha for adults and 0.95 ha for juveniles in palmetto (Sabal spp.) scrub and oak scrub habitats [62]. In Louisiana, winter territories averaged 0.8 to 1.2 ha [57].

COVER REQUIREMENTS:
Red-headed woodpeckers are cavity nesters. The most preferred nest sites are old snags or tree limbs that have been dead for several years, have little or no bark, few large limbs remaining, low canopy cover, and low ground cover [6,20,42,48,49,53,69,75,94]. Nests are most commonly found in dead longleaf pine, loblolly pine, slash pine (P. elliottii), pond pine (P. serotina), plains cottonwood, oaks, maples, birches (Betula spp.), and elms (Ulmus spp.) [5,7,33,35,36,42,48,54,94]. Utility poles and fence posts are used if other nesting sites are limited [43,44,63]. Red-headed woodpeckers typically nest in the largest trees [75] and show a strong preference for medium- to large-diameter nest cavities [15].

In Iowa, 88% of red-headed woodpecker nesting cavities was in snags [83]. Snags with <34% of the total branches remaining were highly favored for nesting. Snags with >66 % of branches remaining were used least [83]. This preference may be related to the softness of the wood due to decay [83,94]. In Wyoming, red-headed woodpeckers nested in limb cavities 80.6% of the time and in boles 19.4% of the time (P<0.001) [36]. Red-headed woodpeckers nested in significantly (P=0.02) more snags (72.2%) than nonsnags (27.8%) [36].

The table below summarizes mean red-headed woodpecker nest site characteristics:

State Habitat type Tree height (m) Tree DBH (cm) Cavity height (m) Diameter of trunk/limb at nest (cm)
Arkansas shortleaf pine-post oak (Q. stellata) 14.8 33.4 11.0 .... [80]
Colorado mature plains cottonwood 15.4 56.9-66.8 8.3-8.4 18.4 [7,75]
Florida longleaf pine-turkey oak (Q. laevis) savanna 13.8 30.6 9.3 .... [94]
Iowa mixed riparian habitats 14.6 .... 9.6 25.7 [84]
Kansas American elm (U. americana)-eastern cottonwood-willow 10.9 .... 7.0 21.8 [48]
Minnesota/Wisconsin closed-canopy mature oak forest 18 54 .... .... [33]
Ohio mixed agricultural and forested habitat or oak-hickory (Carya spp.) stands bordering golf courses 18.3 56.9-58.8 10.2-14.1 20.7 [46,71]
Wyoming mixed cottonwood-willow floodplain, irrigated cropland, and pastureland .... 59.2 10.0 21.2 [36]
Virginia mature oak woodlot 29.5 95.0 17.5 38.0 [14]

Nest cavities in an Illinois study were 23 to 66 feet (7-20 m) in height and always in the bole of a snag [69]. All nest trees in Minnesota and Wisconsin had branch stubs and most had old cavities, <25% bark remaining, and at least one "significant dead portion" [33]. Nest trees in Minnesota and Wisconsin had significantly greater diameters (P<0.001) and were significantly taller (P<0.001) than adjacent trees [33].

Red-headed woodpeckers have strong nest site and nest tree fidelity [5,44]. Cavities may be reused from year to year [5,44,94]. In Ohio, 57.7% of nest cavities had been freshly excavated, while 42.3% were old excavations [46]. No natural cavities were used for nesting in Ohio [46]. In Colorado, both limbs and trunks were utilized nearly equally for nesting, as were old and new cavities [7]. Additionally, nest cavities previously excavated by other species, such as the red-cockaded woodpecker (Picoides borealis) and southern flying squirrel (Glaucomys volans), may be reused by red-headed woodpeckers in subsequent years [20]. Red-headed woodpecker pairs occasionally take over cavities that were occupied by other species earlier in the same breeding season [20]. Rarely, red-headed woodpeckers complete excavation of cavities that were abandoned by other species [43].

Roost trees of nonbreeding individuals in a Florida study were located in longleaf pine and turkey oak snags and in a dead limb of a live turkey oak tree [5]. Juveniles showed a preference for roosting in turkey oaks while adults preferred roosting in longleaf pine snags [5].

FOOD HABITS:
The red-headed woodpecker is omnivorous. Vegetative matter in the diet includes corn (Zea mays), dogwood (Cornus spp.) berries, huckleberries (Gaylussacia spp.), strawberries (Fragaria spp.), blackberries and raspberries (Rubus spp.), mulberries (Morus spp.), elderberries (Sambucus spp.), wild and cultivated black cherries (Prunus serotina), chokecherries (P. virginiana), grapes (Vitis spp.), apples (Malus spp.), pears (Pyrus spp.), pawpaw (Asimina spp.), Carolina lauralcherries (Prunus caroliniana), pecans (Carya illinoensis), acorns, beechnuts, and other seeds [6,35,63,94,96]. Animal foods include beetles (Coleoptera), ants and wasps (Hymenoptera), true bugs (Heteroptera), grasshoppers and crickets (Orthoptera), butterfly and moth larvae (Lepidoptera), spiders (Araneae), myriapods (Myriapoda), mice (Rodentia), small lizards (Iguania), and bird (primarily Passeriformes and Piciformes) eggs and nestlings [6,35,63,94,96]. In winter, red-headed woodpeckers may remain in their summer habitats if sufficient acorn or beechnut mast is available [6]. Maple seeds are an important winter food source for red-headed woodpeckers when other foods are not available [69].

Red-headed woodpeckers exhibit a myriad of behaviors for obtaining and storing food. Perhaps the most conspicuous behavior is flycatching for beetles and other insects [6,48,56,63,91]. Flycatching is less frequent during winter due to a decrease in flying insects during colder months [94]. Red-headed woodpeckers also glean insects, particularly adult beetles, from bark [16,48,94] and feed on insects on the ground [48,69]. Acorns are harvested both directly from trees and from the ground [17,82]. Red-headed woodpeckers may eat acorns immediately after harvesting, or carry them off to a cache [82].

Foods, including acorns and insects, are commonly cached in natural crevices and cavities, which are sealed with wet bark or wood to prevent pillaging by others [6,49]. Insects, such as June beetles (Phyllophaga spp.) and grasshoppers, are cached while still alive and wedged in cracks so they cannot escape [6,35]. Acorns and beechnuts are also stored in cracks and crevices for later use [6].

PREDATORS:
Snakes and mammals commonly predate red-headed woodpecker nests [20]. Northern raccoons (Procyon lotor), eastern racers (Coluber constrictor), and gray ratsnakes (Elaphe obsoleta spiloides) prey on eggs and nestlings [6,44,47]. Flying squirrels (Glaucomys spp.) will consume eggs of the red-headed woodpecker and then take over the nest cavity [3]. Eastern fox squirrels (Sciurus niger) may prey upon eggs and nestlings, although predation by this species has not been directly observed [7,23].

Cooper's hawks (Accipiter cooperi), northern harriers (Circus cyaneus), red-tailed hawks (Buteo jamaicensis), peregrine falcons (Falco peregrinus), eastern screech-owls (Megascops asio), and red foxes (Vulpes vulpes) also prey upon red-headed woodpeckers [27,28,29,35,89].

Nest parasitism by brown-headed cowbirds (Molothrus ater) is extremely rare [31].

MANAGEMENT CONSIDERATIONS:
A steep decline in the global population of red-headed woodpeckers has been documented since 1966 [73]. Nationally, the average decline of the red-headed woodpecker was 2.6% annually between 1966 and 2005. From 1980 to 2005 the average decline was 4.2% annually. The declines for both periods are significant (P<0.01) [73]. In the Midwest, red-headed woodpeckers have declined 3.3% per year for a total decline of 63.8% between 1966 and 1993 [40]. Red-headed woodpecker declines in Florida are of high concern [90].

Collisions with automobiles [6,35] and competition for nesting cavities with other species, particularly European starlings and other woodpeckers, may be contributing to population decline [35,46]. In the past, red-headed woodpeckers were shot because they were considered agricultural pests in fruit groves and cornfields [6,35]. Red-headed woodpeckers also do considerable damage to utility poles and were shot as a result [6].

Habitat loss is likely the primary cause of the decline of the red-headed woodpecker population. Predicted responses of habitat changes on red-headed woodpeckers are [83]:

All woody vegetation removed, resulting in pastures or hayfields Woody vegetation reduced to narrow strips along streams Woody canopy partly removed Woody canopy partly removed and shrubs/saplings thinned Shrubs/saplings thinned Snags removed
Extirpated from community Negative Positive Positive Positive Negative

Snag removal from wooded habitats has a drastic effect on the nesting potential of primary and secondary cavity nesters [83]. Red-headed woodpeckers respond negatively to snag removal [14,21,54,94]. Management recommendations for red-headed woodpeckers in Virginia include retaining snags in woodlots and planting young trees over time to encourage replacement when large trees die [14]. Failing to carry out this recruitment cycle may result in the elimination of mature woodlots in Virginia and the disappearance of the red-headed woodpecker in those areas [14]. Illustrating this point, approximately 53.1% of tree cavities utilized by red-headed woodpeckers were no longer usable after 5 years in a cottonwood bottomland forest in Colorado [76].

In loblolly pine habitat in South Carolina, red-headed woodpeckers were most common where no coarse woody debris was removed (6.8 breeding territories/40 ha) [54]. Red-headed woodpeckers were less common in plots where all down coarse woody debris was removed (3.6 breeding territories/40 ha) and least common in plots where all down coarse woody debris and all standing snags were removed (0.7 breeding territories/40 ha). The differences in breeding territory densities between the 3 stands were significant (P=0.023) [54]. The optimal number of snags required by red-headed woodpeckers in longleaf pine-loblolly pine stands in South Carolina is at least 7 snags/100 acres [39]. At least 1 snag/100 acres is required for red-headed woodpeckers to be present at all in longleaf and loblolly pine stands in South Carolina [39].

Red-headed woodpeckers in Texas were found in clearcuts with standing snags, but not in clearcuts without standing snags [21]. A minimum of 5 snags/ha was assumed adequate for nesting in a clearcut, but a higher density of snags may promote more foraging and nesting on a site. A possible management strategy is to leave large numbers of hardwoods standing during harvesting operations and then killing the standing trees over time to provide a continual recruitment of snags for cavity nesters [21].

Snags should be retained whenever possible [94]. Since tree cavities are ephemeral, recruitment of new snags is crucial for red-headed woodpecker populations to persist.

FIRE EFFECTS AND USE

SPECIES: Melanerpes erythrocephalus

 

  U.S. Department of the Interior, Fish and Wildlife Service, Sherburne National Wildlife Refuge

DIRECT FIRE EFFECTS ON ANIMALS:
The direct effects of fire on red-headed woodpeckers are largely speculative. Fire is unlikely to have major effects on adult birds because adults can fly from smoke and flames [55]. However, fire could have detrimental effects on nestlings and eggs, since their mobility is limited [55].

Only one paper documenting the direct effects of fire on red-headed woodpeckers was found. In the study, a longleaf pine snag containing an active red-headed woodpecker nest ignited during a prescribed May fire in the Florida sandhills [4]. The snag fell on the day of the fire and the nestlings initially survived. The parents continually visited and fed the nestlings. However, by the fourth day, the nestlings had died and were covered with ants [4]. Based on these observations, it is plausible to conclude that the nestlings may have survived if they were within a few days of fledging.

The effects of fire on canopy nesting birds depend primarily on fire severity [55]. Severe surface fires and crown fires may cause injury or death to species nesting in the forest canopy, but such fires typically happen late in the breeding season presumably after many nestlings have fledged. Effects on reproductive success may depend upon the timing of a fire event and any attempts to raise a second brood [55]. See Timing of Major Life History Events for more information on the reproductive biology of the red-headed woodpecker.

Snags are both created and lost during fires. In a red-headed woodpecker study in the sandhills and wet pine flatwoods of central Florida, >33% of nest snags were created during past fires [5]. In other minor habitat types in the study, 83% of nest snags were also created by fire. Prescribed fire later destroyed approximately 50% of the nest snags in the wet pine flatwoods, 70% of nest snags in the sandhills, and no nest snags in the other habitats. The sandhills were dominated by longleaf pine, the flatwoods were dominated by pond pine and longleaf pine, and the other habitats were dominated by sand pine (P. clausa), loblolly pine, and loblolly bay (Gordonia lasianthus) [5].

Fires that destroy nest trees may limit the reproductive potential of canopy-nesting birds. However, taking measures to protect nest trees may minimize the impact fire has on red-headed woodpeckers and their nests. In a review, Robbins and Myers [70] cited a North Carolina study in which red-cockaded woodpeckers were mildly affected by growing-season prescribed fires. Before the fires, flammable materials were cleared from the bases of nest trees. Four red-cockaded woodpecker nests in the study successfully fledged 9 young, while a fifth nest failed. Study recommendations for minimizing deleterious effects of growing-season fire on red-cockaded woodpeckers and their nests include: 1) burning only after nests have been identified; 2) burning only on sites with low fuel accumulation; 3) burning only on days with low ambient temperatures; 4) clearing fuels from the bases of nest trees; and 5) setting backfires to the windward side of nest trees [70]. Since red-headed woodpeckers share some nesting preferences and habitats with the red-cockaded woodpecker, such as the use of large trees for nesting in pine habitats, red-headed woodpeckers may incur similar benefits from prefire site preparation.

Short-term food availability may be improved by fire. Other flycatching birds, such as eastern phoebes (Sayornis phoebe), eastern wood-pewees (Cantopus virens), eastern kingbirds (Tyrannus tyrannus), tree swallows (Tachycineta bicolor), southern rough-winged swallows (Stelgidopteryx ruficollis), and purple martins (Progne subis), actively forage on the multitude of insects available in smoke [51]. Fruits, such as huckleberries and blackberries, that are also utilized as forage may be unavailable during the same year of a burn, but are often abundant during following years [86].

HABITAT-RELATED FIRE EFFECTS:
Oak savanna and other woodland habitats utilized by red-headed woodpeckers historically experienced frequent light- to moderate-severity surface fires [10]. Perhaps relating to an adaptive response to historical fire regimes, red-headed woodpeckers respond favorably toward sites restored through prescribed fire. For instance, red-headed woodpeckers increased in oak savanna restoration sites in Minnesota where prescribed fire was applied every 1 to 2 years over a 32-year period. The increase was likely due to an increase in standing dead trees on the site. The burn units with the highest dead:live tree ratio (0.623:0.865), lowest tree density (74-80 trees/ha), and lowest leaf area index (0.44-0.64) were most suitable to red-headed woodpeckers [19]. In Indiana, red-headed woodpeckers in a 60- to 120-year-old oak-hickory stand were present in prescribed surface fire sites but not in the unburned sites. The fires reduced understory woody vegetation by 40% to 80% and leaf litter by 50% to 80% [2]. Likewise, red-headed woodpeckers in Illinois were more abundant in oak savanna habitats restored through prescribed fire (3.1 individuals/10-point counts) than in closed-canopy forests that had not been burned (0.8 individual\/10-point counts) [9].

In a Kansas tallgrass prairie, red-headed woodpeckers on average were observed in low abundance (0.3 bird/km) in unburned watersheds, while none were observed in burned watersheds. Less woody vegetation was present in the burned sites than in the unburned sites, which may account for the lack of forest-dependent red-headed woodpeckers [100].

In a mature longleaf pine-loblolly pine-shortleaf pine stand with an open hardwood understory, more nonbreeding red-headed woodpeckers were observed in stands burned under prescription during the dormant season (0.16/plot) compared to the number observed in stands burned under prescription during the growing season (0.07/plot) in Georgia. However, the results were not statistically significant (P=0.53). Stands were burned every 3 years [50]. The opposite result was seen in a North Carolina study, where higher densities of red-headed woodpeckers were observed in plots burned under prescription during the growing season than in plots burned under prescription during the dormant season [25].

Mean observations of red-headed woodpeckers were higher in longleaf pine-loblolly pine-shortleaf pine/grassland restoration stands (0.14 detection/point count) than in traditionally managed longleaf pine-loblolly pine-shortleaf pine sawtimber stands (0.06 detection/point count) in Mississippi. The difference was nearly significant (P=0.08). The mixed pine-grassland restoration sites were even-aged stands with rotations ≥70 years, a prescribed burn interval of 2 to 3 years, and mid-story hardwood removal. Traditionally managed mixed pine sawtimber sites in the same study were subject to a 35-year rotation, prescribed burn intervals of 4 to 7 years, and no hardwood removal [98].

Red-headed woodpecker abundance was highest at the beginning of a fire exclusion regime in a loblolly pine-shortleaf pine stand in Florida. The lowest red-headed woodpecker abundance was continuously observed 6 to 15 years after fire exclusion was initiated. Hardwoods had created a thick mid-story up to 16 feet (5 m) following 15 years of fire exclusion [26]. In another Florida study, red-headed woodpeckers were most commonly observed in mature burned sandhills of naturally seeded longleaf pine (>50 years old) where burning had occurred within the last 3 years and at least 1 growing season had passed since the last fire. Red-headed woodpeckers were also regularly observed in mature flatwoods that had been burned within the past 5 years in Florida [90].

The use of silvicultural treatments in addition to fire may alter the postfire response of the red-headed woodpecker. The combined effects of fire and other silvicultural treatments in a Florida longleaf pine habitat were examined. Two years after an initial prescribed burn, experimental treatments of herbicide (hexazinone) and chainsaw felling-girdling of most hardwoods and all sand pine were applied. A burn-only site and a control site were also monitored. A second prescribed burn was conducted in the spring 2 years after the other experimental treatments were applied. The herbicide-treated plots had the lowest abundance of red-headed woodpeckers in the year after the final prescribed burn, but the highest abundance among all treatments a year later. The burn-only plots and the felling-girdling plots had intermediate abundance of red-headed woodpeckers during both years of observation. No red-headed woodpeckers were observed in control plots. None of the differences were significant (P>0.05) [68].

The following table provides fire return intervals for plant communities and ecosystems where the red-headed woodpecker is important. For further information, see the FEIS review of the dominant plant species listed below.

Community or Ecosystem Dominant Species Fire Return Interval Range (years)
maple-beech Acer-Fagus spp. 684-1,385 [13,95]
maple-beech-birch Acer-Fagus-Betula spp. >1,000
silver maple-American elm Acer saccharinum-Ulmus americana <5 to 200
sugar maple Acer saccharum >1,000
sugar maple-basswood Acer saccharum-Tilia americana >1,000 [95]
birch Betula spp. 80-230 [87]
plains grasslands Bouteloua spp. <35 [67,99]
sugarberry-America elm-green ash Celtis laevigata-Ulmus americana-Fraxinus pennsylvanica <35 to 200
beech-sugar maple Fagus spp.-Acer saccharum >1,000
black ash Fraxinus nigra <35 to 200 [95]
green ash Fraxinus pennsylvanica <35 to >300 [24,95]
cedar glades Juniperus virginiana 3-22 [37,67]
yellow-poplar Liriodendron tulipifera <35
shortleaf pine Pinus echinata 2-15
shortleaf pine-oak Pinus echinata-Quercus spp. <10
slash pine Pinus elliottii 3-8
slash pine-hardwood Pinus elliottii-variable <35
sand pine Pinus elliottii var. elliottii 25-45 [95]
South Florida slash pine Pinus elliottii var. densa 1-15 [64,81,95]
longleaf-slash pine Pinus palustris-P. elliottii 1-4 [64,95]
longleaf pine-scrub oak Pinus palustris-Quercus spp. 6-10
pocosin Pinus serotina 3-8
pond pine Pinus serotina 3-8
eastern white pine-northern red oak-red maple Pinus strobus-Quercus rubra-Acer rubrum 35-200
loblolly pine Pinus taeda 3-8
loblolly-shortleaf pine Pinus taeda-P. echinata 10 to <35
Virginia pine Pinus virginiana 10 to <35
Virginia pine-oak Pinus virginiana-Quercus spp. 10 to <35
sycamore-sweetgum-American elm Platanus occidentalis-Liquidambar styraciflua-Ulmus americana <35 to 200 [95]
eastern cottonwood Populus deltoides <35 to 200 [67]
black cherry-sugar maple Prunus serotina-Acer saccharum >1,000
oak-hickory Quercus-Carya spp. <35
northeastern oak-pine Quercus-Pinus spp. 10 to <35 [95]
oak-gum-cypress Quercus-Nyssa-spp.-Taxodium distichum 35 to >200 [64]
southeastern oak-pine Quercus-Pinus spp. <10
white oak-black oak-northern red oak Quercus alba-Q. velutina-Q. rubra <35
northern pin oak Quercus ellipsoidalis <35
bear oak Quercus ilicifolia <35
bur oak Quercus macrocarpa <10 [95]
oak savanna Quercus macrocarpa/Andropogon gerardii-Schizachyrium scoparium 2-14 [67,95]
chestnut oak Quercus prinus 3-8
northern red oak Quercus rubra 10 to <35
post oak-blackjack oak Quercus stellata-Q. marilandica <10
live oak Quercus virginiana 10 to<100 [95]
cabbage palmetto-slash pine Sabal palmetto-Pinus elliottii <10 [64,95]

FIRE USE:
Fires during late spring to mid-summer would likely cause the greatest mortality of nestlings and eggs. Timing prescribed fires outside the height of the nesting season would minimize direct mortality.

Red-headed woodpeckers respond favorably to snag recruitment and retention. Using fire to create new snags and maintain an open habitat would provide nesting sites and facilitate foraging opportunities. Additionally, protecting existing nesting and roosting trees by removing fuels from the bases would minimize the immediate and short-term effects on red-headed woodpecker habitat. If existing snags are destroyed by fire or some other event, recruiting new snags through prescribed fire or mechanical means may allow a red-headed woodpecker population to return to the site in the future.

Melanerpes erythrocephalus: REFERENCES


1. American Ornithologists' Union. 2007. The A.O.U. check-list of North American birds, 7th edition, [Online]. American Ornithologists' Union (Producer). Available: http://www.aou.org/checklist/index.php3. [50863]
2. Aquilani, Steven M.; Morrell, Thomas E.; LeBlanc, David C. 2003. Breeding bird communities in burned and unburned sites in a mature Indiana oak forest. Proceedings of the Indiana Academy of Science. 112(2): 186-191. [60825]
3. Bailey, Harold H. 1913. The birds of Virginia. Lynchburg, VA: J. P. Bell Co. 362 p. [65466]
4. Belson, M. Shane; Small, Parks E. 1998. Uncommon behaviors of red-headed woodpeckers in central Florida. Florida Field Naturalist. 26(2): 44-45. [61941]
5. Belson, Michael Shane. 1998. Red-headed woodpecker (Melanerpes erythrocephalus) use of habitat at Wekiwa Springs State Park, Florida. Orlando, FL: University of Central Florida. 65 p. Thesis. [65156]
6. Bent, Arthur Cleveland. 1939. Life histories of North American woodpeckers. U.S. Natural History Bulletin No. 174. 334 p. [62579]
7. Bergstrom, John T. 1977. Ecology and behavior of woodpeckers in the South Platte River floodplain. Greeley, CO: University of Northern Colorado. 110 p. Dissertation. [65319]
8. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434]
9. Brawn, Jeffrey D. 2006. Effects of restoring oak savannas on bird communities and populations. Conservation Biology. 20(2): 460-469. [61997]
10. Brawn, Jeffrey D.; Robinson, Scott K.; Thompson, Frank R. 2001. The role of disturbance in the ecology and conservation of birds. Annual Review of Ecology and Systematics. 32: 251-276. [55773]
11. Carter, William A. 1967. Ecology of the nesting birds of the McCurtain Game Preserve, Oklahoma. The Wilson Bulletin. 79(3): 259-272. [65476]
12. Clapp, R. B.; Klimkiewicz, M. K.; Futcher, A. G. 1983. Longevity records of North American birds: Columbidae through Paridae. Journal of Field Ornithology. 54: 123-137. [65032]
13. Cleland, David T.; Crow, Thomas R.; Saunders, Sari C.; Dickmann, Donald I.; Maclean, Ann L.; Jordan, James K.; Watson, Richard L.; Sloan, Alyssa M.; Brosofske, Kimberley D. 2004. Characterizing historical and modern fire regimes in Michigan (USA): a landscape ecosystem approach. Landscape Ecology. 19: 311-325. [54326]
14. Conner, Richard N. 1976. Nesting habitat for red-headed woodpeckers in southwestern Virginia. Bird-Banding. 47(1): 40-43. [61966]
15. Conner, Richard N.; Adkisson, Curtis S. 1977. Principal component analysis of woodpecker nesting habitat. The Wilson Bulletin. 89(1): 122-129. [61962]
16. Conner, Richard N.; Jones, Stanley D.; Jones, Gretchen D. 1994. Snag condition and woodpecker foraging ecology in a bottomland hardwood forest. The Wilson Bulletin. 106(2): 242-257. [24214]
17. Cypert, E.; Webster, B. S. 1948. Yield and use by wildlife of acorns of water and willow oaks. Journal of Wildlife Management. 12(3): 227-231. [18951]
18. Daniel, Ryan S.; Fleet, Robert R. 1999. Bird and small mammal communities of four similar-aged forest types of the Caddo Lake area in east Texas. Texas Journal of Science. 51(1): 65-80. [35975]
19. Davis, Mark A.; Peterson, David W.; Reich, Peter B.; Crozier, Michelle; Query, Toby; Mitchell, Eliot; Huntington, Josh; Bazakas, Paul. 2000. Restoring savanna using fire: impact on the breeding bird community. Restoration Ecology. 8(1): 30-40. [35984]
20. Dennis, John V. 1971. Species using red-cockaded woodpecker holes in northeastern South Carolina. Journal of Ornithological Investigation. 42(2): 79-87. [26097]
21. Dickson, James G.; Conner, Richard N.; Williamson, J. Howard. 1983. Snag retention increases bird use of a clear-cut. Journal of Wildlife Management. 47(3): 799-804. [13855]
22. Dickson, James G.; Noble, Robert E. 1978. Vertical distribution of birds in a Louisiana bottomland hardwood forest. The Wilson Bulletin. 90(1): 19-30. [60732]
23. Doherty, Paul F., Jr.; Grubb, Thomas C., Jr.; Bronson, C. L. 1996. Territories and caching-related behavior of red-headed woodpeckers wintering in a beech grove. The Wilson Bulletin. 108(4): 740-747. [61930]
24. Eggler, Willis A. 1980. Live oak. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 63-64. [49984]
25. Engstrom, R. T.; McNair, D. B.; Brennan, L. A.; Hardy, C. L.; Burger, L. W. 1996. Influence on birds of dormant versus lightning-season prescribed fire in longleaf pine forests: experimental design and preliminary results. In: Wadsworth, Kelly G.; McCabe, Richard E., eds. Facing realities in resource management: Transactions of the 61st North American wildlife and natural resource conference; 1996 March 22-27; Tulsa, OK. 61. [Place of publication unknown]: [Publisher unknown]: 200-207. [62434]
26. Engstrom, R. Todd; Crawford, Robert L.; Baker, W. Wilson. 1984. Breeding bird populations in relation to changing forest structure following fire exclusion: a 15-year study. The Wilson Bulletin. 96(3): 437-450. [60735]
27. Errington, Paul L. 1933. Food habits of southern Wisconsin raptors. Part II. Hawks. The Condor. 35(1): 19-29. [61995]
28. Errington, Paul L. 1937. Food habits of Iowa red foxes during a drought summer. Ecology. 18(1): 53-61. [61985]
29. Errington, Paul L.; Breckenridge, W. J. 1936. Food habits of marsh hawks in the glaciated prairie region of north-central United States. The American Midland Naturalist. 17(5): 831-848. [61986]
30. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]
31. Friedmann, Herbert. 1963. Hosts of the brown-headed cowbird. In: Host relations of the parasitic cowbirds. United States National Museum Bulletin No. 233. Washington, DC: United States National Museum: 41-172. [61104]
32. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998]
33. Giese, Collette L. Adkins; Cuthbert, Francesca J. 2005. Woodpecker nest tree characteristics in upper midwestern oak forests. The Canadian-Field Naturalist. 119(3): 367-376. [65353]
34. Graber, Jean W.; Graber, Richard R. 1979. Severe winter weather and bird populations in southern Illinois. The Wilson Bulletin. 91(1): 88-103. [65135]
35. Graber, Jean W.; Graber, Richard R.; Kirk, Ethelyn L. 1977. Illinois birds: Picidae. Biological Notes No. 102. Urbana, IL: State of Illinois, Department of Registration and Education, Natural History Survey Division. 73 p. [64983]
36. Gutzwiller, Kevin J.; Anderson, Stanley H. 1987. Multiscale associations between cavity-nesting birds and features of Wyoming streamside woodlands. The Condor. 89(3): 534-548. [65355]
37. Guyette, Richard; McGinnes, E. A., Jr. 1982. Fire history of an Ozark glade in Missouri. Transactions, Missouri Academy of Science. 16: 85-93. [5170]
38. Hardin, Kimberly I.; Evans, Keith E. 1977. Cavity nesting bird habitat in the oak-hickory forests--a review. Gen. Tech. Rep. NC-30. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 23 p. [13859]
39. Harlow, Richard F.; Guynn, David C., Jr. 1983. Snag densities in managed stands of the South Carolina coastal plain. Southern Journal of Applied Forestry. 7(4): 224-229. [12571]
40. Herkert, James R. 1995. An analysis of midwestern breeding bird population trends: 1966-1993. The American Midland Naturalist. 134(1): 41-50. [26795]
41. 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]
42. Ingold, Danny J. 1989. Nesting phenology and competition for sites among red-headed and red-bellied woodpeckers and European starlings. The Auk. 106: 209-217. [61938]
43. Ingold, Danny J. 1990. Simultaneous use of nest trees by breeding red-headed and red-bellied woodpeckers and European starlings. The Condor. 92(1): 252-253. [61936]
44. Ingold, Danny J. 1991. Nest-site fidelity in red-headed and red-bellied woodpeckers. The Wilson Bulletin. 103(1): 118-122. [61934]
45. Ingold, Danny J. 1994. Influence of nest-site competition between European starlings and woodpeckers. The Wilson Bulletin. 106(2): 227-241. [24096]
46. Ingold, Danny J. 1994. Nest-site characteristics of red-bellied and red-headed woodpeckers and northern flickers in East-Central Ohio. Ohio Journal of Science. 94(1): 2-7. [62016]
47. Jackson, J. A. 1969. Observations at a nest of the red-headed woodpecker. Museum of Natural History, annual report, 1968-1969: 3-10. [65134]
48. Jackson, Jerome A. 1976. A comparison of some aspects of the breeding ecology of red-headed and red-bellied woodpeckers in Kansas. The Condor. 78(1): 67-76. [61965]
49. Kilham, Lawrence. 1983. Red-headed woodpecker. In: Paynter, Raymond A., Jr., ed. Life history studies of woodpeckers of eastern North America. Publications of the Nuttall Ornithological Club, No. 20. Cambridge, MA: Nuttall Ornithological Club: 113-128. [65711]
50. King, T. Gregory; Howell, Mark A.; Chapman, Brian R.; Miller, Karl V.; Schorr, Robert A. 1998. Comparisons of wintering bird communities in mature pine stands managed by prescribed burning. The Wilson Bulletin. 110(4): 570-574. [36037]
51. Komarek, E. V., Sr. 1969. Fire and animal behavior. In: Proceedings, annual Tall Timbers fire ecology conference; 1969 April 10-11; Tallahassee, FL. No. 9. Tallahassee, FL: Tall Timbers Research Station: 161-207. [13531]
52. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. [3455]
53. Kurta, Allen; Kath, Joseph; Smith, Eric L.; Foster, Rodney; Orick, Michael W.; Ross, Ronald. 1993. A maternity roost of the endangered Indiana bat (Myotis sodalis) in an unshaded, hollow, sycamore tree (Platanus occidentalis). The American Midland Naturalist. 130(2): 405-407. [53799]
54. Lohr, Steven M.; Gauthreaux, Sidney A.; Kilgo, John C. 2002. Importance of coarse woody debris to avian communities in loblolly pine forests. Conservation Biology. 16(3): 767-777. [60758]
55. Lyon, L. Jack; Telfer, Edmund S.; Schreiner, David Scott. 2000. Direct effects of fire and animal responses. In: Smith, Jane Kapler, ed. Wildland fire in ecosystems: Effects of fire on fauna. Gen. Tech. Rep. RMRS-GTR-42-vol. 1. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 17-23. [44435]
56. MacRoberts, Michael H. 1970. Notes on the food habits and food defense of the acorn woodpecker. The Condor. 72: 196-204. [8117]
57. MacRoberts, Michael H. 1975. Food storage and winter territory in red-headed woodpeckers in northwestern Louisiana. The Auk. 92(2): 382-385. [61967]
58. Martin, Thomas E. 1980. Diversity and abundance of spring migratory birds using habitat islands on the Great Plains. The Condor. 82(4): 430-439. [61992]
59. Martin, Thomas E. 1995. Avian life history evolution in relation to nest sites, nest predation, and food. Ecological Monographs. 65(1): 101-127. [61994]
60. Moorman, Christopher E.; Guynn, David C., Jr. 2001. Effects of group-selection opening size on breeding bird habitat use in a bottomland forest. Ecological Applications. 11(6): 1680-1691. [43241]
61. Morse, Douglas A. 1970. Ecological aspects of some mixed-species foraging flocks of birds. Ecological Monographs. 40(1): 119-168. [65352]
62. Moskovits, Debra. 1978. Winter territorial and foraging behavior of red-headed woodpeckers in Florida. Wilson Bulletin. 90(4): 521-535. [61957]
63. Mumford, Russell E.; Keller, Charles E. 1984. The birds of Indiana. Bloomington, IN: Indiana University Press. 376 p. [60761]
64. Myers, Ronald L. 2000. Fire in tropical and subtropical ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 161-173. [36985]
65. National Geographic Society. 1999. Field guide to the birds of North America. 3rd ed. Washington, DC: The National Geographic Society. 480 p. [60563]
66. Niemi, Gerald J. 1978. Breeding birds of burned and unburned areas in northern Minnesota. Loon. 50: 73-84. [14451]
67. Paysen, Timothy E.; Ansley, R. James; Brown, James K.; Gottfried, Gerald J.; Haase, Sally M.; Harrington, Michael G.; Narog, Marcia G.; Sackett, Stephen S.; Wilson, Ruth C. 2000. Fire in western shrubland, woodland, and grassland ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 121-159. [36978]
68. Provencher, Louis; Gobris, Nancy M.; Brennan, Leonard A.; Gordon, Doria R.; Hardesty, Jeffrey L. 2002. Breeding bird response to midstory hardwood reduction in Florida sandhill longleaf pine forests. Journal of Wildlife Management. 66(3): 641-661. [42243]
69. Reller, Ann Willbern. 1972. Aspects of behavioral ecology of red-headed and red-bellied woodpeckers. The American Midland Naturalist. 88(2): 270-290. [61970]
70. Robbins, Louise E.; Myers, Ronald L. 1992. Seasonal effects of prescribed burning in Florida: a review. Misc. Publ. No. 8. Tallahassee, FL: Tall Timbers Research, Inc. 96 p. [21094]
71. Rodewald, Paul G.; Santiago, Melissa J.; Rodewald, Amanda D. 2005. Habitat use of breeding red-headed woodpeckers on golf courses in Ohio. Wildlife Society Bulletin. 33(2): 448-453. [61908]
72. Rumble, Mark A.; Gobeille, John E. 2004. Avian use of successional cottonwood (Populus deltoides) woodlands along the middle Missouri River. The American Midland Naturalist. 152: 165-177. [1217]
73. Sauer, J. R.; Hines, J. E.; Fallon, J. 2005. Red-headed woodpecker Melanerpes erythrocephalus: North American breeding bird survey trend results. In: The North American breeding bird survey, results and analysis 1966-2005. Version 6.2.2006, [Online]. U.S. Department of the Interior, Geological Survey, Patuxent Wildlife Research Center, Migratory Bird Research (Producer). Available: http://www.mbr-pwrc.usgs.gov/bbs/bbs.html [2007, January 29]. [65354]
74. Sedgwick, James A.; Knopf, Fritz L. 1986. Cavity-nesting birds and the cavity-tree resource in plains cottonwood bottomlands. Journal of Wildlife Management. 50(2): 247-252. [19447]
75. Sedgwick, James A.; Knopf, Fritz L. 1990. Habitat relationships and nest site characteristics of cavity-nesting birds in cottonwood floodplains. Journal of Wildlife Management. 54(1): 112-124. [11105]
76. Sedgwick, James A.; Knopf, Fritz L. 1992. Cavity turnover and equilibrium cavity densities in a cottonwood bottomland. Journal of Wildlife Management. 56(3): 477-484. [19280]
77. Sharps, Jon C.; Uresk, Daniel W. 1990. Ecological review of black-tailed prairie dogs and associated species in western South Dakota. The Great Basin Naturalist. 50(4): 339-344. [14895]
78. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. [23362]
79. Sibley, Charles G.; Monroe, Burt L., Jr. 1990. Distribution and taxonomy of the birds of the world. New Haven, CT: Yale University Press. 1111 p. [22814]
80. Smith, Kimberly G.; Withgott, James H.; Rodewald, Paul G. 2000. Red-headed woodpecker--Melanerpes erythrocephalus. In: Poole, A.; Gill, F., eds. The birds of North America. No. 518. Philadelphia, PA: The Academy of Natural Sciences; Washington, DC: The American Ornithologists' Union: 1-27. [61928]
81. Snyder, James R.; Herndon, Alan; Robertson, William B., Jr. 1990. South Florida rockland. In: Myers, Ronald L.; Ewel, John J., eds. Ecosystems of Florida. Orlando, FL: University of Central Florida Press: 230-274. [17391]
82. Sork, Victoria L.; Stacey, Peter; Averett, John E. 1983. Utilization of red oak acorns in non-bumper crop year. Oecologia. 59: 49-53. [4593]
83. Stauffer, Dean F.; Best, Louis B. 1980. Habitat selection by birds of riparian communities: evaluation effects of habitat alterations. Journal of Wildlife Management. 44(1): 1-15. [8118]
84. Stauffer, Dean F.; Best, Louis B. 1982. Nest-site selection by cavity-nesting birds of riparian habitats in Iowa. The Wilson Bulletin. 94(3): 329-337. [62017]
85. Stevenson, Henry M.; Anderson, Bruce H. 1994. The birdlife of Florida. Gainesville, FL: University of Florida Press. 892 p. [60776]
86. Stoddard, Herbert L., Sr. 1963. Bird habitat and fire. In: Proceedings, 2nd annual Tall Timbers fire ecology conference; 1963 March 14-15; Tallahassee, FL. Tallahassee, FL: Tall Timbers Research Station: 163-175. [18997]
87. Swain, Albert M. 1978. Environmental changes during the past 2000 years in north-central Wisconsin: analysis of pollen, charcoal, and seeds from varved lake sediments. Quaternary Research. 10: 55-68. [6968]
88. Tappe, Philip A.; Thill, Ronald E.; Peitz, David G.; Perry, Roger W. 2004. Early succession bird communities of group-selection openings and clearcuts in the Ouachita Mountains, Arkansas and Oklahoma. In: Guldin, James M., tech. comp. Ouachita and Ozark Mountains symposium: ecosystem management research; [Dates & location unknown] Gen. Tech. Rep. SRS-74. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station: 42-54. [55740]
89. Toland, Brian R. 1990. Nesting ecology of red-tailed hawks in central Missouri. Transactions, Missouri Academy of Science. 24: 1-16. [22703]
90. Tucker, James W.; Hill, Geoffrey E.; Holler, Nicholas R. 2003. Longleaf pine restoration: implications for landscape-level effects on bird communities in the lower Gulf Coastal Plain. Southern Journal of Applied Forestry. 27(2): 107-121. [44563]
91. Twomey, Arthur C. 1945. The bird population of an elm-maple forest with special reference to aspection, territorialism, and coactions. Ecological Monographs. 15(2): 173-205. [65511]
92. U.S. Department of the Interior, Fish and Wildlife Service, Division of Endangered Species. 2007. Threatened and endangered animals and plants, [Online]. Available: http://www.fws.gov/endangered/wildlife.html [2007, February 22]. [62042]
93. Van Dyke, Fred; Van Kley, Sarah E.; Page, Christy E.; Van Beek, Jodi G. 2004. Restoration efforts for plant and bird communities in tallgrass prairies using prescribed burning and mowing. Restoration Ecology. 12(4): 575-585. [54784]
94. Venables, Ann,; Collopy, Michael W. 1989. Seasonal foraging and habitat requirements of red-headed woodpeckers in north-central Florida. Tallahassee, FL: Florida Game and Fresh Water Fish Commission, Nongame Wildlife Program. 49 p. [61978]
95. Wade, Dale D.; Brock, Brent L.; Brose, Patrick H.; Grace, James B.; Hoch, Greg A.; Patterson, William A., III. 2000. Fire in eastern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 53-96. [36983]
96. Wiebe, Karen L.; Koenig, Walter D.; Martin, Kathy. 2006. Evolution of clutch size in cavity-excavating birds: the nest site limitation hypothesis revisited. The American Naturalist. 167(3): 343-353. [61902]
97. Willson, Mary F. 1970. Foraging behavior of some winter birds of deciduous woods. The Condor. 72(2): 169-174. [61918]
98. Wood, Douglas R.; Burger, L. Wes, Jr.; Bowman, Jacob L., Hardy, Carol L. 2004. Avian community response to pine-grassland restoration. Wildlife Society Bulletin. 32(3): 819-829. [61603]
99. Wright, Henry A.; Bailey, Arthur W. 1982. Fire ecology: United States and southern Canada. New York: John Wiley & Sons. 501 p. [2620]
100. Zimmerman, John L. 1992. Density-dependent factors affecting the avian diversity of the tallgrass prairie community. The Wilson Bulletin. 104(1): 85-94. [60823]

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