The Red-cockaded woodpecker is found in the Southeastern United States. Populations are distributed as far west as Oklahoma and Texas, north to Kentucky, south into Florida and east to the Alantic Ocean. Primarily concentrated in certain old-growth pine forests. (Poole et al., 1994; Winkler et al., 1995)

Biogeographic Regions: nearctic (Native )

endemic to a single nation

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

Global Range: (200,000-2,500,000 square km (about 80,000-1,000,000 square miles)) Historical range included the southeastern Piedmont and Coastal Plain from New Jersey to Texas, and inland to Kentucky, Tennessee, Missouri, and Oklahoma (AOU 1983, Jackson 1994). Now the species is virtually extirpated north of North Carolina and in all interior states except Arkansas (Walters 1991). Populations are fragmented and most are quite small (Stangel et al. 1992). The six largest populations are in the Apalachicola National Forest (Florida), North Carolina Sandhills, Francis Marion National Forest (South Carolina), Kisatchie National Forest (Louisiana), Eglin Air Force Base and Blackwater State Forest (Florida), and Red Hills hunting plantations in southern Georgia (James 1995).

Length: 20 to 23 cm Small, white and black woodpecker with a barred back. This characteristic distinguishes it from all other species of the same genus within its range. Large white feathers overlie the ears and cheek area. Underside white or grayish, with notable black spots along the sides of the breast. Male has red spots located on each side of the nape that are seldomly exposed. Female is slightly larger and lack red spots.

(Poole et al., 1994; Short, 1982)

Average mass: 56 g.

Length: 18 cm

Weight: 44 grams

• Terrestrial

These woodpeckers are adapted to mature, living, open-pine forests that are frequently maintained by naturally occurring summer fires. They are particularly dependent on a few species of pinewith substantially strong heartwood. Older trees (70 years or more) are preferred. Species utilized include longleaf (Pinus palustris), slash (P. elliottii), shortleaf (P. echinata), pond (P. serotina) and pitch (P. rigada) pines. (Poole et al., 1994; Short, 1982; Winkler et al., 1995)

Terrestrial Biomes: forest

Comments: Habitat consists of open, mature pine woodlands, rarely deciduous or mixed pine-hardwoods located near pine woodlands (Steirly 1957, Hooper et al. 1980, U.S. Fish and Wildlife Service 1980, Kalisz and Boettcher 1991). Optimal habitat is characterized as a broad savanna with a scattered overstory of large pines and a dense groundcover containing a diversity of grass, forb, and shrub species (Hooper et al. 1980, AOU 1991). Midstory vegetation is sparse or absent (Hooper et al. 1980, Locke et al. 1983, Hooper et al. 1991, Loeb et al. 1993).

The open, park-like characteristic of the habitat is maintained by low intensity fires, which occurred historically during the growing season at intervals of about 1-10 years (Christensen 1981, SNN 1990), evidenced by the rapid invasion of hardwoods into pine savannas following fire exclusion for brief periods (less than 10 years), the high annual frequency of lightning strikes in these habitats in summer (Platt 1988), and the pyrogenic nature of characteristic plant species such as longleaf pine (Pinus palustris) (Rebertus et al. 1989) and wiregrass (Aristida stricta) (Platt et al. 1988). AOU (1991) concluded that this bird "...is a symbol of fire-maintained, old-growth pine savannas, once the dominant ecosystem in the Southeast."

CAVITY TREES

Nesting and roosting occur in tree cavities. Active cavity trees are almost exclusively old, living, flat-topped pine trees (but see Patterson and Robertson 1983, Hooper 1982). Typical noncavity roosts include scars on the trunks of trees, forks on live pines (Hooper and Lennartz 1983), and the underside of large limbs (Baker, pers. comm.). However, juveniles often investigate dead trees and may roost in them until they obtain a cavity in a living tree (Ligon 1970).

Excavation through the sap wood appears to limit the speed with which cavities are constructed, perhaps because sap leaking into the tunnel interrupts excavation (Hooper et al. 1980, Walters 1991). The tunnel is excavated back into the tree at an upward slope such that resin or pitch from the sapwood drains from the hole, and that rain water is prevented from collecting in the cavity (Ligon et al. 1980, Jackson 1978b). Even so, Teulings (1973) reported that many nest cavities were flooded with rain following violent storms.

Once birds have tunneled through the sapwood and into the heartwood, they excavate downward forming a gourd-shaped chamber 15-25 cm deep and 1.7- 12.7 cm wide. The girth of the tree must be wide enough to allow the central chamber to be excavated in the resin-less heartwood, hence one of the reasons for selecting old trees (Ligon et al. 1980).

Roosting and nesting cavities have been found in longleaf, loblolly (Pinus taeda), shortleaf (Pinus echinata), slash (Pinus ellioti), pond pine (Pinus rigida), and even bald cypress (Taxodium disthicus) (Dennis 1971). Some evidence suggests longleaf pine is preferred even when mature stands of other pine species are available (Hopkins and Lynn 1971, Lay and Sweptson 1973, Baker 1981, Lennartz et al. 1983, Hovis and Labisky 1985, Ligon et al. 1986). The historic distribution of longleaf coincides with the region where Audubon (1839) reported the greatest abundance of Red-cockaded Woodpeckers. In addition, relict stands of old-growth longleaf today have some of the highest densities of this species (Engstrom 1982, Carter et al. 1983).

Even so, other species such as shortleaf and loblolly pine are important in areas where longleaf pine is uncommon or absent (Jackson 1971, Wood 1983a, Shapiro 1983).

Red-cockaded woodpeckers show a clear preference for older trees (Jackson et al. 1979, Jackson and Jackson 1986, Delotelle and Epting 1988, Engstrom and Evans 1990), contrary to conclusions reached by Field and Williams (1985). The average age of cavity trees ranges from 63-126 years for longleaf pine, 70-90 years for loblolly pine, 75-149 years for shortleaf pine, 62-130 years for pond pine, and >70 years for slash pine (Hooper et al. 1980).

The diameter of cavity trees at breast height (dbh) is usually at least 35 cm (Lay and Russel 1970, Jones and Ott 1973, Hovis and Labisky 1985), except in south Florida where cavity trees averaged only about 25 cm dbh (Shapiro 1983, Delotelle et al. 1983, Beaver and Dryden 1992) and may be as small as 15.4 cm dbh (Beaver and Dryden 1992).

Cavity trees are generally infected with red heart disease, a fungus (Phellinus pini) that attacks the heartwood, destroys cell walls, and causes the wood to become soft and pithy (Steirly 1957). Conner and Locke (1982) also found cavities in wood infected by one of four other fungi species. Although red heart disease is apparently not a strict prerequisite for cavity excavation (Conner and Locke 1982, Beckett 1971, Jackson 1977), Hooper et al. (1991) suggested there is a universal preference for trees with decayed heartwood. An independent group of researchers agreed with this statement (SNN 1990).

In addition to requirements for old pine trees, appropriate habitat also includes open, park-like conditions extending across the area surrounding a cluster of cavity trees Walters (1991).

Shapiro (1983) found that the average minimum distance between cavity trees was 104 m in south Florida, whereas Wood (1983) reported an average minimum distance of only 58 m in Oklahoma.

Various measurements of the vegetative structure surrounding clusters of cavities have been compared with different types of control sites lacking woodpeckers. However, the fidelity this species shows to cavity sites (Harlow 1983, Nesbitt et al. 1983a, Lennartz et al. 1987, Walters 1991), coupled with social interactions (Walters 1990), mean that measurements around "active cavities" may not indicate productive habitat (Hooper et al. 1980, Walters 1990, Walters 1991); a pair of birds or a solitary male, and even some nesting groups may remain at a cavity associated with low-quality habitat for many years (Walters 1989, 1991).

The basal area of pine overstory is one of the more important habitat components within cavity clusters. Hooper et al. (1980) maintained that appropriate conditions require a mature pine stand (more than 70 years old) with a stocking density of 11.5-18 sq m of overstory pines per ha. This stocking density is relatively high compared to that recommended by others. Conner and O'Halloran (1987) recommended a pine basal area of 9-14 sq m per ha, while Hovis and Labisky (1985) recommended about 12 sq m per ha. The Recovery Plan (Lennartz and Henry 1985) recommended that cavity clusters have a pine basal area of 5.8-8.4 sq m per ha. The basal area of pines reported from studies in south and central Florida is much less than suggested by Hooper et al. (1980), averaging 4.0-7.0 sq m per ha (Delotelle et al. 1983, Shapiro 1983, Nesbitt et al. 1983b).

Multivariate techniques have been used to quantify habitat conditions surrounding active cavity clusters. Locke et al. (1983) measured 29 vegetative variables in a 0.04 ha area around active cavity trees and compared these with measurements at randomly selected old pines. Four variables were correlated with active sites: 1) the total number of hardwood trees (6.4 in occupied sites versus 12.8 in random sites); 2) the number of understory pines (6.9 in occupied sites versus 3.5 in control); 3) the mean ratio of bole length to height of pines greater than 28 cm diameter breast height (0.48 in occupied sites versus 0.43 in random sites); and 4) the number of midstory hardwoods (1.1 in occupied sites versus 1.8 in random sites).

FORAGING HABITAT

Less is known about foraging habitat than about conditions needed around cavity clusters. Hooper et al. (1980) stated that "... the best cavity site is no good if adequate foraging habitat is unavailable." Recent studies (Delotelle et al. 1983, Hovis and Labisky 1985, Conner and Rudolph 1991b, Loeb et al. 1992) and anecdotal evidence provide some general indication of the importance of foraging habitat. However, this species appears to use a wide range of pine and hardwood habitats (Hooper et al. 1980, U.S. Fish and Wildlife Service 1985), and foraging behavior is influenced by conspecifics, group size, and many other factors (Conner and Rudolph 1991b, Walters 1991). These conditions make it difficult to ascertain with certainty the importance of specific habitat features within foraging areas.

Several lines of evidence point to a problem with large cleared tracts. Walters (pers. comm.) monitored a population of 12 cavity clusters for 9 years and found the population to be stable until logging cleared much of the foraging habitat in the area. Soon afterwards, the population showed signs of falling apart. Hooper et al. (1980) reported that sites were abandoned where extensive clear cuts were created within the foraging area. They also proposed that, when clearing reduces the foraging habitat to less than 40 ha, persisting groups may have difficulty raising young. Conner and Rudolph (1991b) found that the removal of forest cover within 800 m of cavity clusters was associated with cluster inactivation. The effects of forest removal were particularly noticeable in small populations.

Foraging occurs in a diversity of forested habitat types that includes pines of various ages as well as some hardwood-dominated habitats. Despite this seemingly catholic use of different habitat types, most foraging appears to take place on older pine trees or in open pine habitats (Baker and Thompson 1971, Hooper et al. 1980, Hooper and Lennartz 1981, Delotelle 1983).

Delotelle et al. (1983) found that live pine stems greater than 23 cm dbh represented only 19% of available foraging substrate in central Florida but received 65% of the use, and also found that longleaf pine was used as the foraging substrate 90% of the time.

Similarly, Porter and Labisky (1986) found that longleaf pines made up only 40% of their study area yet were the foraging substrate approximately 77% of the time.

As described elsewhere, females forage on trunks more often than do males, and an abundance of trees with large trunk surfaces may be more important to females; Jackson (pers. comm.) suggested that females lost weight after pines larger than 25 cm dbh were removed from all but a small buffer surrounding cavity trees..

Other evidence supporting the need for foraging habitat consisting of large areas of old pines lies in the fact that the largest populations occur where old-growth pine trees and low hardwood densities extend throughout a broad area (Engstrom 1982, Hooper et al. 1980, Delotelle et al. 1983, Conner and Rudolph 1991a).

The Recovery Plan (Lennartz and Henry 1985) suggested suitable foraging habitat requires 6350 stems > 25.4 cm diameter at breast height, and 796 sq m basal area of pine stems in stands >30 years old within 1.3 km (0.5 miles) of a cavity cluster. These values do not apply in south Florida (Beaver and Dryden 1992) where birds use larger areas.

LANDSCAPE HABITAT FEATURES

Landscape features, such as fragmentation of foraging habitat, total area of foraging habitat, percentage of pinewood or hardwood cover, contiguity of the canopy and forest cover, and habitat patch size and shape may affect the habitat quality (Hooper et al. 1980, SNN 1990, Conner and Rudolph 1991b). The importance of such variables is not well known (Walters 1991), but a growing body of research focuses on this issue, particularly on some public lands where timber harvest patterns may create unfavorable landscapes (Conner and Rudolph 1991b). A potential problem in such research is the key role that cavity trees play in determining whether an area is ever actually used by red- cockaded woodpeckers (Walters 1991). Areas that have suitable habitat characteristics, yet lack suitable cavity trees, will not likely be occupied by red-cockaded woodpeckers (Walters 1991), and thus some comparisons will be misleading.

Seagle et al. (1992) compared characteristics of forest compartments with active colonies and those having no colonies. Active clusters were associated with: 1) an increased acreage of mature longleaf pine; 2) an increased acreage of all pine species; 3) a decreased percentage of acres of longleaf, loblolly, and slash pines in stands less than 20 acres in size; 4) a decreased percentage of acres of mature loblolly pine; and 5) a decreased acreage of loblolly pine between ages 20 and 39 years.

Conner and Rudolph (1991b) found that foraging habitat could be fragmented and isolated as a result of forest-harvest patterns, and that larger groups of woodpeckers had consistently fewer clear cuts near cavity sites. Fragmentation did not appear to have an effect on dispersal (e.g., the ability of dispersing females to find unmated males), but it did apparently affect the quality of foraging habitat. Conner and Rudolph (1991b) warned that it may be possible to have a sufficient quantity of foraging habitat within 800 m of an active cluster but still have insufficient arrangement of foraging habitat. Fragmentation influenced a group's access to foraging habitat by forcing birds to go through territories of adjacent groups. This increases the probability of cluster inactivation.

Non-Migrant: Yes. At least some populations of this species do not make significant seasonal migrations. Juvenile dispersal is not considered a migration.

Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

Picoides borealis is omnivorous, and its diet consists mainly of adult, larvae and eggs of tree surface and subsurface arthropods, especially beetle larvae and ants. Usually forages for food on pine trees by ripping loose bark from the surface with an upward or sideways movement of the bill or in some cases, bark is stripped away with the feet. To a lesser extent, various seeds, nuts (pecans) and fruit are consumed, and occasionally the woodpecker frequents cornfields in search of earworms in a certain larval stage that are residing within the corn kernel. (Bent, 1992; Poole et al., 1994; Short, 1982; Winkler et al., 1995)

Comments: Locate food primarily by sight and by exploring cavities, crevices, and tunnels with the long tongue (Jackson 1983a). Apparently do not use sound to locate prey (Jackson 1983a), as do some other woodpeckers (Callegari 1955). While searching for food, the bill and/or feet are used to help pry off pieces of bark and expose prey. Most foraging time is spent scaling bark from living pines. (Murphrey 1939). One foraging technique involves backing down the tree and flaking off bark with the feet while catching prey with the bill (U.S. Fish and Wildlife Service 1980). Murphrey (1939) described the primary foraging technique as a spiral ascent of the trunk. Visits to lightning-struck trees infested with beetle larvae are common (Hooper et al. 1980). Occasionally they will also "fly catch" prey using an aerial foraging technique similar to that of Red-headed Woodpeckers (Beckett 1971).

An analysis of 99 stomachs revealed a diet of 84% insects and 16% berries (Beal 1911). Insects eaten are primarily ants, spiders, cockroaches, centipedes, termites, and the larvae and eggs of various other insects. More recently, C. Hess (pers. comm.) found that a species of wood-boring ant (Crematogaster SP.) constitutes more than 80% of food items fed to young birds on the Apalachicola National Forest. Hess suggested that an understanding of the biology of this ant may be important in management decisions. In South Carolina, wood roaches (Parcoblatta sp.) were the dominant prey fed to nestlings; other common nestling foods were wood boring beetle larvae, Lepidoptera larvae, spiders, and ants (Hanula and Franzreb 1995).

Especially gregarious when foraging (Murphrey 1939). May also associate with flocks of Brown-headed Nuthatches (Jackson 1983c) or eastern bluebirds (Beckett 1971), especially in winter (Murphrey 1939).

Males and females exhibit a divergence in foraging behavior (Ligon 1970, Skorupa 1979, Ramey 1980, Hooper and Lennartz 1981). Males tend to forage on limbs of the crown and midtrunk more frequently than do females, while females tend to forage more frequently on the lower trunk; sexes show considerable overlap on the mid-trunk (Porter et al. 1985). In one study, the average foraging height of males was 14.1 m and that of females was only 8.7 m. The mean difference in foraging height ranges from about 3.7 to 5.4 m (Hooper and Lennartz 1981, Porter et al. 1985); this difference greatest January-March (7.4 m, Hooper and Lennartz 1981), while both sexes tend to use the canopy during breeding season (SNN 1990). Sexes used trees of similar size and species, and the methods for capturing prey were also similar (Lennartz and Hooper 1981). Adults bring food to the nest from as far away as 640 m (Ligon 1970).

Birds drink water from flooded holes in trees and from the ground (Murphrey 1939, Hooper et al. 1980).

Note: For many non-migratory species, occurrences are roughly equivalent to populations.

Estimated Number of Occurrences: 81 to >300

Comments: This species is represented by a large number of occurrences (subpopulations), but approximately 50% of all known birds occur in just six populations (James 1995), while the remaining birds are scattered across >130 sites.

James (1995) presented the following 1990 state-by-state estimates for the number of active "cavity clusters:" Alabama 157; Arkansas 132; Florida 1224; Georgia 596; Kentucky 4; Louisiana 464; Mississippi 154; North Carolina 465; South Carolina 615; Oklahoma 15; Tennessee 1; Texas 315; Virginia 5.

10,000 - 100,000 individuals

Comments: USFWS (2003) reported that there are an estimated 14,068 red-cockaded woodpeckers living in 5,627 known active clusters across eleven states. USFWS (http://www.fws.gov/rcwrecovery/rcw.html, August 2007 version) reported the population as consisting of 6,000 groups or 16,000 birds. U.S. Department of Defense and USFWS (2006) reported the populaton size as 15,150 adults. Only the public lands component of these estimates are based on recent and rigorous survey information (Florida Fish and Wildlife Conservation Commission 2001).

Only three populations number more than 300 breeding groups (Walters 1991, James 1995).

TERRITORIALITY AND HOME RANGE: Maintain territories throughout the year, and appear to recognize precise boundaries (Ligon 1970). Groups seem to be least cohesive during the incubation period and when nestlings are being fed (Walters 1990). Helpers assist in the defense of territories (Ligon 1970), and territorial displays include characteristic vocalizations (Ligon 1970), wing-spreading displays, wing flattering, raising of the red cockades (in the case of males, Hagan and Reed 1988), and physical attack (Walters 1990). Drumming is not as common a territorial display in red-cockaded woodpeckers as it is in other woodpeckers, but it may occur when territorial birds encounter an alien bird (Hooper et al. 1982).

Walters (1990) reported that adult males are the primary defenders against intruder males, and adult females against intruder females, for most of the year. During the breeding season, however, both sexes respond to a single intruder. Birds are also most aggressive towards intruders and neighboring groups during the breeding season. Outside the breeding season, a resident group may sometimes forage peaceably with an intruder within the territory, or near a neighboring group along a territory boundary.

A comprehensive 3-year study by Hooper et al. (1982) reported home ranges from 34-225 ha (mean of total ranges 86.9 ha, mean of year-round ranges 70.3 ha). In other studies, home range estimates varied from 15-220 ha and averaged around 67 ha (Baker 1977, Crosby 1971, Skorupa and McFarlane 1976, Nesbitt et al. 1978, Sherrill and Case 1980, Nesbitt et al. 1983b, Wood 1983a), although extremely large territories of about 400 ha may exist (Hooper et al. 1980).

Home range estimates may include "extra-territorial" areas that are utilized by other neighboring conspecifics as well as floaters in the population (Walters 1989); these are not strongly defended and are estimated to average about 8.4 ha (Hooper et al. 1982), but may be as large as 30 ha (Hooper et al. 1982, Repasky 1984, Blue 1985, Porter and Labisky 1986, DeLotelle et al. 1987). Repasky (1984) suggested that extra-territorial range is underestimated because sampling is usually not extensive in winter and late summer when birds use larger areas. However, in the few studies in which home range size estimates were subdivided into territorial and extra-territorial areas, similar mean territory sizes of 70 ha were obtained (Hooper et al. 1982, Repasky 1984, Blue 1985).

Seasonal variation in range size has been reported, though debate on this issue exists. Skorupa and McFarlane (1976) found that home ranges were smallest in summer and largest in winter. These authors warned that average range values may be misleading since "... the increased winter foraging requirements of the species must be considered ..." as part of conservation and management efforts. Wood (1983) also found home range to be larger in winter in Oklahoma (52.8 ha) than in summer (44.1 ha) and spring (26.0 ha). However, these data were for a single, 5-member clan. In their larger study, Hooper et al. (1982) found larger (although not statistically significant) home ranges in spring (mean 49 ha) and summer (mean 47.7 ha) than in fall (mean 37.5 ha). They further found that birds had larger home ranges during the post-fledging period (mean 43.1 ha) than during the nestling period (mean 27.8). Jerauld et al. (1983) noted a restricted range during the first few days following fledging. They suggested that range sizes decreased during this time when fledglings had difficulty following adults.

Range size also appears to vary geographically to some degree, with larger home ranges occurring in south and central Florida. An early study of a single group in south Florida reported a range of 159 ha (Patterson and Robertson 1981). In central Florida, DeLotelle et al. (1983) found that four year-round range were 120-203 ha and averaged 150 ha. Nesbitt et al. (1983b) reported that five groups in southwest Florida had an average home range size of 148 ha and a range of 80-218 ha. Nesbitt et al. (1983b) noted that this species probably requires larger ranges at the southern limits of their distribution owing to poor habitat quality.

DISPERSAL: Although dispersal is primarily undertaken by young birds; mate loss and an apparent avoidance of inbreeding sometimes cause adults to disperse, and adults may also occasionally move to neighboring territories for unknown reasons (Walters 1989). In North Carolina, fledgling females dispersed an average of 4.8 km, maximum 31.5 km; breeding males dispersed an average of 2.1 km, maximum 15 km; fledgling males dispersed an average of 5.1 km, maximum 21.1 km; helper males dispersed an average of 1.8 km, maximum 17.1 km; and solitary males dispersed an average of 2.3 km, maximum 8.5 km (Walters 1989).

The relatively short dispersal distance implies that rates of inbreeding may be high even though close inbreeding is avoided (Walters 1990). That is, matings between second cousins may be common while parent-offspring matings are avoided. This may have led to the high similarities of DNA profiles reported by Haig et al. (1993b). However, Walters (1988) described a long-distance dispersal event for one female, which moved 90 km and seemed to follow a highway corridor that contained appropriate habitat conditions. The bird also traversed unfavorable habitats.

SURVIVAL CHARACTERISTICS: Reed et al. (1988) and Walters (1989) reported that the average annual rate of survival for birds in their first year is 57% for males and 32% for females. Females experience lower survival rates primarily as a result of the increased frequency of dispersal in this sex. Annual survival rates for 2-3-year-old birds is relatively high: approximately 77% for males and 75% for females. About 25% of fledgling males live to be at least 4 years old, while only about 13% of fledgling females live to this age. Fewer than 1% of fledgling birds live to be 10 years old.

Major storms are one of the primary sources of mortality. Hurricane Hugo killed about 63% of the birds found on the Francis Marion National Forest (Hamrick 1992). Accipiter hawks may prey on adults (Ligon 1970). When an accipiter flies into an area, an alarm call is given and all birds "freeze" with their bills pointing straight up (Ligon 1970). They will often wait in this manner ten minutes after the hawk has left before resuming foraging (Ligon 1970). Other sources of mortality for adults include being stuck in resin (Locke et al. 1979) and killed by rat snakes while incubating eggs (J. Kappes, pers. comm.). No disease outbreaks have been reported.

EFFECTS ON OTHER SPECIES: At least 25 vertebrates (Kappes undated) have been recorded using red-cockaded woodpecker cavities. As many as 56% of the cavities in a cluster may be used by other species (Harlow and Lennartz 1983). Yellow-bellied sapsuckers have been observed feeding at resin wells (Rudolph et al. 1991). The sticky resin associated with cavity trees may pose an occasional hazard to woodpeckers and other birds (Locke et al. 1979, Barnett et al. 1983, Locke et al. 1979). Jackson (1983c) reported that brown-headed nuthatches seemed to follow foraging red-cockaded woodpeckers and gleaned anything the woodpeckers dislodged but did not consume. Beckett (1971) described similar behavior for the eastern bluebird. Morse (1970) also observed brown-headed nuthatches foraging in association with red-cockaded woodpeckers in Louisiana but felt that the woodpeckers were following the nuthatches.

Average lifespan

Status: wild: 193 months.

Picoides borealis is monogamous and pair bonding takes place throughout the year, whenever a female arrives in an unoccupied cavity cluster or nesting site. They partake in cooperative breeding, and a pair may or may not have helpers, which are usually male. Males are able to breed at one year, but breeding is usually delayed because young males are serving as helpers in their own natal group. Copulation can be seen at any time, increasing in occurrence in late spring. The first brood is laid between April and June, and a second brood in the same season is rare. The clutch contains 2-5 eggs. Incubation is done by both parents and usuallly begins after the second egg is laid, with the breeding male remaining on the nest throughout the night. Incubation lasts 10-13 days. Both parents feed the nestlings and development is rapid. Most chicks fledge after 26-29 days, but commonly remain dependent on the parents for up to 5 months. The nest is located in the roost cavity of the breeding male. These cavities are excavated in mature pines between 12-100 feet up. They are 8-12 inches deep, with a 2 inch diameter entrance that tilts upward to prevent pitch and water from entering the nest.

(Poole et al., 1994; Short, 1982; Winkler et al., 1995)

Average time to hatching: 12 days.

Average eggs per season: 3.

Average age at sexual or reproductive maturity (male)

Sex: male: 240 days.

Average age at sexual or reproductive maturity (female)

Sex: female: 240 days.

COOPERATIVE BREEDING SYSTEM: Live in groups containing a single breeding pair and zero to four 'helpers' (Walters 1990), rarely as many as nine (Hooper et al. 1980). Helpers help incubate eggs, feed nestlings and fledglings, and defend territories (Ligon 1970, Lennartz and Harlow 1979, Walters 1990). Mated pairs are monogamous (Ligon 1970, Walters 1990, Haig et al. 1993b). Walters et al. (1988) reported that male-female pairs were the most common social unit (60%); about 30% of the groups contained one or more adult helpers and about 10% consisted of solitary males; only 5% of groups had more than one helper.

Each member of a group usually has an exclusive roost cavity (Hooper and Lennartz 1983b, Harris and Jerauld 1983, Jansen 1983). Access to a cavity is critical to the nesting success of males, since the nesting cavity is almost always the cavity of the single breeding male (Ligon 1970, Hooper and Lennartz 1983).

It takes many months to excavate a cavity (Hooper et al. 1980, Walters 1991). The importance of attaining a cavity, contrasted with the extended time required to excavate one, has led (in part) to different strategies among young birds for coping with the common situation wherein most suitable cavities are occupied by conspecifics (Walters 1990). Almost all young females and most young males disperse and find an existing cavity with a new group (Walters et al. 1988). Another strategy, employed by 27 per cent of the young males, is to remain on the natal territory in hopes of inheriting the territory or another nearby territory. (Walters et al. 1988). Only very rarely do young birds disperse to new areas and excavate new cavities (Walters 1990).

Birds that remain in natal territories may do so for many years and help the breeding pair raise and care for new birds (Walters et al. 1988).

Once a male attains breeding status in a group, it usually retains that position until death. Females may switch groups after attaining breeding status, particularly when an offspring male inherits a territory; this may help avoid close inbreeding (Walters et al. 1989).

NESTING: Copulation observed November-December and March-May (Ligon 1970, Crosby 1971, Baker 1971), but egg laying usually occurs April-early May. Groups may not nest every year (Hopkins and Lynn 1971), perhaps in response to environmental factors.

Clutch size 2-5, 3-4 usual (Murphrey 1939). Incubation lasts about 10-12 days (Hooper et al. 1980), and often begins with the laying of the second egg. This means that some eggs hatch earlier than others, giving early hatchlings a size and competitive advantage over other nestlings; Ligon (1970) reported that later-hatching young often starved within 24 hours.

Nestlings are altricial and remain in the nest for 26-29 days (Ligon 1970). Ligon (1970) reported hatching rates of about 90% but fledgling rates of only 50%.

Neal et al. (1993a) found that roughly half of the older nestlings in their study area were eaten by snakes. Red-bellied Woodpeckers have also been observed pulling nestlings out of nests (Ligon 1971), and flying squirrels (Glaucomys volans) may destroy nests (Walters 1991). Bad weather represents another major source of nestling mortality (Neal et al. 1993b).

Lennartz et al. (1987) found that breeding groups with helpers fledge more young per year (2.05) than groups consisting of a single breeding pair (1.40).

Reproductive output increases with age, especially after approximately 2-3 years (Reed et al. 1988, Walters 1989).

Gowaty and Lennartz (1985) reported that the sex ratios of nestlings and fledglings favored males 59:41. This bias suggests that female nestlings may be more difficult to raise or males somehow have an advantage during the nestling period. Nests tended by a breeding pair and helpers produced more even sex ratios than did nests tended by a single pair (Gowaty and Lennartz 1985).

FLEDGING: Young birds leave the nest after 26-29 days (Ligon 1970), and most nesting activity is finished by early July (Baker 1983a, Wood 1983b). Within 3-5 days, fledglings are able to follow adults on extended foraging trips (Ligon 1971). Juveniles may beg for and receive food from adults for 5 months or longer (Ligon 1970).

Juveniles progress through a series of molts before attaining adult plumage; molting may become particularly heavy towards late summer and food requirements increase during these periods of heavy molt (Jackson 1983a). Post-juvenile plumage is attained by late fall or early winter of the first year (Jackson 1983a).

• 2008
  Vulnerable
• 2007
  Vulnerable
• 2004
  Vulnerable
• 2000
  Vulnerable
• 1996
  Vulnerable
• 1994
  Vulnerable

Due to its dependence on particular southern pine forests for food and habitat, the Red-cockaded woodpecker has been considered endangered since 1968. Habitat fragmentation, clear-cutting, midstory encroachment of hardwood and natural disasters, including Hurricane Hugo in 1989, have been the major causes of the devastating decline in this species. Remaining populations are concentrated on federal and private lands with the largest located on state land in Florida. There have been two recovery plans written to conserve and rebuild the populations of the woodpecker. One was established in 1979, but was never fully acted upon. Then, in 1985 the U.S. Fish and Wildlife Service introduced a slightly revised plan. It was based heavily on research by Fish and Wildlife Service biologists on the then largest, healthiest population, located in the Francis Marion National Forest. This plan omitted some sugestions from the previous 1979 plan, including the idea of linking fragmented populations with corridors made from interstate highways, and added others. The general objective of the revised plan was to ultimately achieve range-wide recovery. A specific objective objective was to establish a viable population that consisted of a minimum effective population size of 500 breeding birds. This would maintain desired levels of genetic variation for long-term population survival. The potential carrying capacity was calculated for different areas. Each carrying capacity depended on species compostion, structure and age of forest habitats available. The approximate overall capacity ranged from one clan per 400 acres of habitat to one clan per 200-250 acres of habitat. The American Ornithologist Union highly criticised the efforts of the Fish and Wildlife Service. One of the reported problems was their estimation of the species minimum viable population size. Some argued that the estimation should be 1,018 breeding birds, which in turn, would increase the habitat to 25,450 ha of pine forest, a much larger calculation than suggested by the revised plan. Despite the arguments, this plan has not been changed and no new plan has been written.

Recently, however, new approaches to conservation including old cavity restoration, artificial cavity construction, and the introduction of females into isolated groups, have made some positive advancments towards the increase in populations. The total population is now estimated at about 7,500 individuals.

(Poole et al., 1994; Winkler et al., 1995; U.S. Fish and Wildlife Service, 1985)

IUCN Red List of Threatened Species: vulnerable

United States

Rounded National Status Rank: N3 - Vulnerable

Rounded Global Status Rank: G3 - Vulnerable

Reasons: Fairly large range in the southeastern United States, but both quantity and quality of suitable habitat are much reduced; historical extent of suitable habitat and probably population size have been reduced by about 97 percent. Short-term rotation timber management eliminated mature diseased pines required for roosting, nesting, and foraging; fire suppression allowed invasion of pine stands by hardwoods. Recent management innovations (e.g., prescribed burns, cavity management) have alleviated threats and resulted in population increases in most areas, but a stable or increasing trend independent of continuing artificial cavity installation (a short-term solution) can be achieved only when large old pines are available in abundance. Further population increases, independent from continuing artificial cavity installation eventually should allow the rank to be changed to G4 (apparently secure).

Intrinsic Vulnerability: Highly vulnerable

Comments: Red-cockaded woodpeckers do not exhibit any of the various types of metapopulation structure (see USFWS 2003). Local extinction followed by natural recolonization from another population is extremely unlikely for this species (USFWS 2003).

Environmental Specificity: Very narrow. Specialist or community with key requirements scarce.

Other Considerations: As of October 2007, all states within the range of this species ranked it as SX, SH, S1, or S2 (extirpated, historical, critically imperiled, or imperiled).

Global Short Term Trend: Relatively stable (=10% change)

Comments: In the 1990s, in response to intensive management (prescribed burning, cavity management) based on a new understanding of population dynamics and new management tools, most populations were stabilized and many showed increases. Other populations remain in decline, and most have small population sizes (USFWS 2003). Most core populations within recovery units are located in national forests, and most populations in national forests appear to be stable or increasing; a few are in decline (USFWS 2003).

PRESENT TRENDS: Most of the species' population lives in Florida, and the Florida Fish and Wildlife Conservation Commission (2001) estimated that populations there have increased 20 per cent since 1990. However, some of this increase is the result of increased inventory effort, not actual population increases, and the Florida Ornithological Society (2001) countered that statewide trends cannot be inferred from present data.

FUTURE TRENDS: The Florida Fish and Wildlife Conservation Commission (2001) predicted that most, if not all occurrences on private land there could disappear within the next twenty years, and there would be a total decline in the Florida population of greater than 20 per cent over the same time. However, the Florida Ornithological Society (2001) pointed out that these predictions do not take into account the infrequent but potentially catastrophic effects of hurricanes on local populations; when they included these effects into a predictive model, they found that the vulnerability of individual populations increased tenfold.

Global Long Term Trend: Decline of >90%

Comments: Populations have declined by more than 97 per cent over the past 100 years (USFWS 2000, James 1995, Noss 1989). The species is extirpated in New Jersey, Maryland, Tennessee, Missouri and Kentucky. Early naturalists reported the species to be very abundant throughout its range (Audubon 1839, Howell 1932). Howell (1932) listed the red-cockaded woodpecker as "...the most abundant woodpecker..." in one north-central Florida county.

Despite protection under the U.S. Endangered Species Act, all monitored populations except one declined in size between 1970 and the early 1980s (U.S. Fish and Wildlife Service 2000). James (1995) conservatively estimated that the range-wide population declined by at least 23% between the early 1980s and 1990.

The population on the Francis Marion National Forest (South Carolina) was thought to be increasing until Hurricane Hugo destroyed most of the cavity trees and 80 per cent of the foraging habitat; the population declined about 80 per cent (Hooper et al. 1990, Hooper and McAdie 1995).

Reports from many smaller populations also indicate a general decline. DeFazzio et al. (1987) and Haig et al. (1993a) described a declining population on the Savannah River Site. Baker (1983a) reported on the extirpation of a population in north Florida. A severe decline was recorded in the 1980s in eastern Texas, due largely to hardwood invasion of midstory and to isolation of cavity clusters (Conner and Rudolph 1991a, Conner and Rudolph 1991b). However, the Piedmont-Hitchiti population in the Georgia Piedmont remained relatively stable from the early 1980s to the early 1990s (Stevens 1992). In Oklahoma, the number of groups and number of individuals in a 3795-ha area declined by 62% and 75%, respectively, between 1977 and 1989-1990 (Kelly et al. 1994). See Costa and Escano (1989), McFarlane (1992), and James (1995) for information on the trends of other populations.

See Eddleman and Clawson (1987) for information on the status of habitat in Missouri.

Degree of Threat: A : Very threatened throughout its range communities directly exploited or their composition and structure irreversibly threatened by man-made forces, including exotic species

Comments: Threatened by a loss of habitat (either gradually through poor management or rapidly through the outright destruction of old-growth forests), forest fragmentation, competition with other species for cavities, catastrophic events, and demographic and genetic processes affecting populations confined to isolated conservation areas (U.S. Fish and Wildlife Service 1985, Ligon et al. 1986, Walters 1991). These threats do not exist independent of one another, and collectively they may render even large populations susceptible to extinction processes over a narrow window of time (Ligon et al. 1986, Walters 1991, SNN 1990).

The dependence of this species on old-growth pine forest is the single most critical factor leading to its endangered status (AOU 1991). This habitat requirement is in direct conflict with timber management policies on some public and almost all private lands (Jackson 1986, Ligon et al. 1986, AOU 1991). Private timber stands in the southeastern U.S. are generally on short rotations (less than 45 years) that do not permit trees to attain the characteristics sought by red-cockaded woodpeckers (Neel 1971, Ligon et al. 1986, Jackson 1976). Overall, only 2.5% of the current pine acreage in the southeastern U.S. is considered suitable nesting habitat (U.S. Fish and Wildlife Service 1985), and most of this exists on public lands. The few stands of old-growth timber remaining on private lands are under increasing pressures to be converted to short-rotation pine plantations (Neel 1971), and legal provisions for maintaining habitat on private lands are weak (Ligon et al. 1986).

Management on many public lands focuses on maintenance of "middle-aged" trees rather than old-growth forests (Jackson 1986, Ligon et al. 1986), which again is contrary to the biological requirements of this species. Hovis and Labisky (1985), for example, found that 92% of the cavity trees on the Apalachicola National Forest were more than 80 years old, yet more than 50% of the pines on this National Forest were less than 60 years old. The consequences of such management can be seen in downward population trends reported for most major populations on public lands (Ligon et al. 1986, AOU 1991, Walters 1991) as well as the potentially unstable conditions reported for some larger populations (James 1991). Since red-cockaded woodpeckers require mature forests and have limited capacities to colonize new areas (Walters 1991), errors in management decisions may take a long time to correct. A potential decline in the growth of southeastern pine forests (Zeide 1992) could further exacerbate problems on public lands. A stable or increasing trend independent of continuing artificial cavity installation (a short-term solution) can be achieved only when large old pines are available in abundance (USFWS 2003).

Hardwood encroachment is also a persistent problem on some public lands owing to the infrequent use of prescribed fires (Hooper et al. 1980, U.S. Fish and Wildlife Service 1980, Jackson 1986, Ligon et al. 1986, Walters 1991). Hardwood encroachment has been implicated in the decline of numerous populations (Walters 1991). Beckett (1971) and Crosby (1971) were among the first to suggest that red-cockaded woodpeckers abandoned cavity trees and clusters if the hardwood midstory reached the height of the cavity entrance. This has generally been confirmed in work by Thompson and Baker (1971), Carter (1974), Van Balen and Doerr (1978), Wood (1983), and Loeb et al. (1992). It is believed that hardwood trees clustered around cavity trees provide gray rat snakes or flying squirrels, a potential nest predator and a potential nest usurper (Jackson 1974, Jackson 1978b), with access to cavities without having to cross fresh resin (Dennis 1971, Jackson 1974). Hardwood encroachment may also increase interspecific competition for cavities (Costa and Escano 1989). Finally, hardwood encroachment may also affect the flight path to a cavity (Wood 1983a, Kelly et al. 1993) or the quality of foraging habitat (Conner and Rudolph 1991a).

Conner and Rudolph (1991b) found that fragmentation and isolation created by forest-harvest patterns also may threaten some populations. Since few large populations exist on lands that are not managed also for timber production (James in press), fragmentation resulting from timber harvest may threaten many populations. In comparisons of cavities with single males versus cavity clusters with breeders and helpers, Conner and Rudolph (1991b) found the larger groups had fewer clear cuts near cavity sites and less fragmentation of the available foraging habitat (see above). Conner and Rudolph (1991b) warn that it is possible to have a sufficient quantity of foraging habitat within 800 m of an active cluster, but still have insufficient arrangement of foraging habitat. In addition, Conner et al. (1991) suggest that the pattern of clear cuts could increase mortality of cavity trees through wind damage.

Competition for existing cavities is a pernicious problem and may threaten some small populations (Jackson 1978a, Harlow and Lennartz 1983, Rudolph et al. 1990b, Loeb 1993). Baker (1983) monitored the decline and eventual extirpation of a small red-cockaded woodpecker population in north Florida and found that competition for cavities and aggressive interactions between red-cockaded woodpeckers and other species seemed to increase in frequency and intensity during the decline. The southern flying squirrel, which may occupy 10-21% of the cavities in some areas (Loeb 1993), is perhaps the most common usurper of red-cockaded cavities (Jackson 1978b, Harlow and Lennartz 1983, Rudolph et al. 1990, Loeb 1993), followed by (in approximate order of frequency of use) red-bellied woodpecker, red-headed woodpecker, eastern bluebird, northern flicker, great crested flycatcher (Myiarchus crinitus), and tufted titmouse (Parus bicolor) (Jackson 1978b, Harlow and Lennartz 1983, Rudolph et al. 1990). Red-bellied woodpeckers may be the most frequent nest usurpers in some areas (Ligon 1970). Other species observed using woodpecker cavities very infrequently include gray rat snake, brown-headed nuthatch, broad-headed skink (Eumeces laticeps), gray tree frog (Hyla versicolor), fox squirrel (Sciurus niger), screech owl (OTIS Asio), five-lined skink (Eumeces fasciatus), and honey bee (Apis mellifera) (Baker 1971, Dennis 1971b, Jackson 1978, Harlow and Lennartz 1983). Data on the invertebrates using red-cockaded woodpecker cavities is rarely collected (Dennis 1971a, b, Baker 1971). Usurpation of cavities by any of these species may be to the detriment of red-cockaded woodpeckers. Flying squirrels were the major competitor for nest cavities in Texas (Rudolph et al. 1990) and South Carolina (Loeb 1993), but competition was not thought to be a major factor influencing the stability of the woodpecker population in Texas (Rudolph et al. 1990). There is little evidence of predation by flying squirrels during the nesting season (Harlow and Doyle 1990). Walters (1991) contended that flying squirrels represent a minor management problem since they do not enlarge the entrance of the cavity. On the other hand, other types of woodpeckers may significantly modify cavities, and once cavity has been modified, it is rarely used again by red- cockaded woodpeckers (Walters 1991). The pileated woodpecker is particularly destructive since it enlarges a large number of red-cockaded cavities (Jackson 1978a, Walters 1991). There are at least two reasons why cavities enlarged by other woodpeckers are abandoned. Enlarged cavities frequently fill with rain water, and enlarged cavities enable avian and mammalian predators to remove roosting red-cockaded woodpeckers (Jackson 1978).

Conner and Rudolph (1995) documented a high rate of southern pine beetle-caused mortality of cavity trees in the Angelina National Forest in Texas. This resulted in a high rate of use of artificial cavities. Trees with cavity inserts may produce less resin and provide less favorable protection against snake predation.

CATASTROPHIC EVENTS: Hurricanes, epizootic diseases, and, to a lesser extent, beetle infestations) affect populations periodically. The isolated nature of existing populations makes catastrophic events a cause for concern since natural recolonization is unlikely. When Hurricane Hugo passed through South Carolina on September 21- 22, 1989, it destroyed almost half of the 100 km2 of mature forest on the Francis Marion National Forest (Hooper et al. 1990). The storm reduced the population of woodpeckers by a staggering 63%, and it destroyed 87% of known active cavity clusters (Hooper et al. 1990). Although about 700 birds survived, there were only 225 cavities remaining. The hurricane also destroyed an estimated 50-60% of the foraging habitat, and it is believed that it may take 75 years for the forest to be suitable habitat for red-cockaded woodpeckers (Hooper et al. 1990, Hamrick 1992). Cavity trees may also be killed by the southern pine beetle (Berlanger et al. 1988, Conner et al. 1991). In Texas, pine beetles killed 53% of the 453 cavity trees monitored (Conner et al. 1991).

FRAGMENTATION: The fragmentation and isolation of managed populations within the historic range of the species may threaten to reduce genetic diversity (Ligon et al. 1986) and increase the probabilities of extinction as a result of demographic and environmental fluctuations (Shaffer 1981, Walters 1991). Stangel et al. (1992) found that genetic heterozygosity at 16 presumed gene loci was weakly correlated with population size (smaller populations having less heterozygosity). However, heterozygosity in all populations analyzed fell within "normal" ranges for birds, and this correlation appeared to be heavily influenced by data from two small populations. Ligon et al. (1986) stressed that threats posed by inbreeding and genetic deterioration could extinguish small populations, while Haig et al. (1993a) proposed that inbreeding may present a serious to the viability of small populations (<20 active clusters).

Walters (1991) cautioned against the use of genetic criteria in assessing the importance of populations. The genetic models used to predict population viability are extremely imprecise (Koenig 1988, Reed et al. 1993), and analyses presented by Lande (1988) suggested that demography and environmental stochasticity pose a much greater threat to small populations. Stangel et al. (1992) also cautioned against treating small populations as "lost causes" based on the perceived threat of genetic deterioration.

Restoration Potential: The time required for restoration depends mostly on the age of potential cavity trees in an area, the degree of hardwood encroachment, the size of the area (i.e., the capacity to support several active cavity clusters), and other conditions. If suitable habitat conditions do not exist, restoration may take many decades. Hooper et al. (1990) estimated that it may take 75 years for the Francis Marion National Forest to become suitable red- cockaded woodpecker habitat following the effects of Hurricane Hugo.

The slow rate of habitat colonization exhibited by red-cockaded woodpeckers (Walters 1990) implies it will be difficult to re-establish extirpated populations. Population augmentation is being used to enhance populations on some national forests (Hess and Costa, in press). However, a reintroduction attempted in an unoccupied area in Georgia (Odom 1983) did not result in the establishment of a population. After 2.5 years, only 2 of the 12 translocated birds were known to survive, although one pair was observed breeding in the first year following translocation (Odom 1983). Other translocation programs have met with mixed success (Reinman 1984, DeFazio et al. 1987).

Preserve Selection and Design Considerations: The minimum amount of land needed for a single group of birds likely falls within the general range of 40-160 hectares (Hooper et al. 1980, SNN 1990); habitat quality will influence the specific area requirements (Conner and Rudolph 1991a). South Florida range sizes lie towards the upper end of the range provided here; adjustments should be made accordingly.

The area needed to sustain a "viable" population will depend on the definition of viability. SNN (1990) suggested that sites with fewer than 50 active clusters were vulnerable to extinction. SSN (1990) furthermore suggested the following thresholds can be used as a coarse gauge of population stability, with stability increasing dramatically at each level:

Level 1. An unoccupied area, but with potential habitat that could be occupied, should be a minimum of 40-160 hectares.

Level 2. A population of 6-7 breeding pairs that can be sustained for a 20-year period, or longer, even in an isolated condition, would require about 240-1120 hectares.

Level 3. A population with a "genetically desirable" condition of 25 active clusters capable of short-term viability (e.g. 50+ years), would require a minimum area of about 1,000-4,000 hectares.

Level 4. A "desirable condition" of 400 breeding pairs to achieve long-term recovery of populations would require a minimum area of about 16,000-64,000 hectares.

An additional consideration (SNN 1990, Walters 1991) is the percentage of solitary males in the population. This percentage varies from site to site (10-35%) (Walters 1990, James 1991), and minimum area requirements may need to be increased by 10-35% accordingly. However, appropriate habitat and population management should be capable of bringing area requirements towards the lower end of the ranges described above, except perhaps in South Florida where consistently larger home ranges are reported (Nesbitt et al. 1983a, Delotelle et al. 1983).

Another preserve design consideration is the imperative for frequent prescribed burning. Large-scale burning may be difficult in preserves near dense human populations, airports, and major highways. See Brenner and Wade (1992) for information on "smoke easements" in areas where residential developments adjoin management areas (Brenner and Wade 1992).

The short dispersal distances characteristic of this species imply that preserve designs should focus on populations as isolated groups (SNN 1990). The chances of populations frequently interacting over a large distances are slim (Simberloff et al. 1992), and efforts to connect some populations may prove ineffective. SNN (1990) recommended that 27 km represented the maximum distance over which two semi-isolated populations might interact frequently. SNN (1990) warned that a stable population would not likely consist of isolated clusters strung out towards the outer extreme of this estimated dispersal distance. See Jackson (1976), Walters (1988), and Ligon et al. (1986) for discussion of the possible use of highway rights-of-way as dispersal corridors.

According to Lennartz and Henry (1985) and SSN 1(1990), 15 distinct populations, each consisting of 500 active clusters, are needed to assure the future survival of the species. Dispersion of such populations probably will be dictated largely by the distribution of existing large public lands, but data on geographic genetic variation (Stangel et al. 1992) suggest that preserves should include populations from across the range of the species.

A broad geographic dispersion of protected populations also reduces the threat posed by somewhat localized events such as hurricanes (Shaffer 1981, Cox et al. 1994). The protection or enhancement of non-coastal populations may also help to guard against hurricanes.

Management Requirements: See ES.OPT1 and ES.OPT2 for MGMT.REQS.

Management Programs: U.S. Forest Service: Robert G. Hooper (Southeastern Forest Experiment Station, Charleston, SC): involved with the use of artificial cavity structures to augment/restore populations. David Allen (U.S. Forest Service, Southeastern Forest Experiment Station, Department of Forest Resources, Clemson University, SC): involved with translocation experiments on national forests. Dennis Krusac and Jim Walker (U.S. Forest Service, Atlanta, GA): analyzing the effects of new timber-harvest regimes. Forrest L. Oliveria (Forest Pest Management, Pineville, LA): studying pine beetle management. Warren G. Montague (Poteau Ranger District, Fayetteville, AK): studying means of reducing mortality to snake predation and cavity losses to squirrel activities. Ernest E. Stevens (Department of Forest Resources, Clemson University, Clemson, SC): analyzing population viability. Craig Watson (Forest Service, Moncks Corner, SC): involved with the restoration of woodpecker population on the Francis Marion National Forest.

U.S. Department of Defense: Carl Petrick (Natural Resource Branch, Eglin Air Force Base, Niceville, FL) and Bob Progulske (Avon Park Air Force Base, Avon Park, FL): investigating new burning programs and developing models for ecosystem management. Stephen Parris (Environmental Division, Fort Polk Air Force Base, Fort Polk, LA): monitoring the impacts of a new multi-purpose range complex.

U.S. Fish and Wildlife Service: Ralph Costa (Red-cockaded Woodpecker Coordinator, U.S. Fish and Wildlife Service, College of Forestry and Recreation Resources, Clemson University, Clemson, SC): involved with habitat conservation planning and the review of agency management programs. Joe Reinman (St. Marks National Wildlife Refuge, St. Marks, FL) and David Richardson (Noxubee National Wildlife Refuge, Brooksville, MS): applied intensive management techniques to 2 small populations.

University Personnel: David Kulhavy (College of Forestry, Stephen F. Austin State University, Nacogdoches, TX): working on the impact of southern pine beetles, the effects of midstory removal on cavity clusters, and landscape disturbances in cavity clusters. Joseph Neal (Arkansas Cooperative Fish and Wildlife Service, University of Arkansas, Fayetteville, AK): developing exclusion devises for rat snakes and flying squirrels. J. H. Carter, III, and Jeffrey Walters (Department of Zoology, North Carolina State University, Raleigh, NC): investigating the use and effectiveness of artificial cavity structures. Richard N. Conner and D. Craig Rudolph (Southern Forest Experiment Station, Stephen Austin State University, Nacogdoches, TX): working on dynamics of cavity excavation and effects of wind damage. Jerome Jackson (Department of Biological Sciences, Mississippi State University, Mississippi State, MS): investigating the effects of timber harvest.

State Agencies: Robert Epting (St. Johns River Water Management District, Palatka, FL): worked on forest stand selection and management. Jim Cox (Florida Game and Fresh Water Fish Commission, Tallahassee, Florida): worked on conservation planning and application of GIS technology.

Nongovernmental Groups: R. Todd Engstrom (Tall Timbers Research Station): using artificial cavities and developing conservation strategies for private lands in south Georgia. Reed Bowman (Archbold Biological Station): measuring response of populations to different management programs. Roy DeLotelle (DeLotelle and Guthrie, Inc., Gainesville, FL): developed management plans for a population on private lands in central Florida.

A regional conservation program is being attempted for private hunting plantations in south Georgia (Anonymous, no date). Most of these lands have been profitably managed using a selective harvest regime for nearly 50 years (Neel 1971).

Monitoring Programs: U.S. Forest Service: Robert G. Hooper (Southeastern Forest Experiment Station, Charleston, SC): determining suitable monitoring programs for the U.S. Forest Service, monitoring impacts of hurricanes. Ron Escano (U.S. Forest Service, Atlanta, GA): monitoring population recovery and forest management.

University Personnel: Susan Loeb and Kay Franzreb (Southeastern Forest Experiment Station, Department of forest Resources, Clemson University): respectively, monitoring turnover rates of nest cavities and the use of global positioning systems. J. H. Carter, III, and Jeffrey Walters (Department of Zoology, North Carolina State University, Raleigh, North Carolina): monitor a large population using banding techniques and annual cavity searches. Frances James (Florida State University, Tallahassee, Florida): applied vegetation and population monitoring techniques to the population on the Apalachicola National Forest, monitoring the dynamics of cavity use in 2 populations in Florida.

U.S. Department of Defense: Carl Petrick (Natural Resource Branch, Eglin Air Force Base, Niceville, FL) and Bob Progulske (Avon Park Air Force Base, Avon Park, FL): mapped cavity locations using geographic information systems technology and developed state-of-the-art monitoring procedures. Similarly detailed programs have been completed for Avon Park Air Force Range (Bob Progulske, Avon Park Air Force Range, Avon Park, FL) and Ft. Benning (John Duvesky, The Nature Conservancy, Ft. Benning, GA).

State Agencies: Brent Ortego (Texas Parks and Wildlife Department, Austin, TX): investigating monitoring nesting success in Texas national forests. Dana Bradshaw (Virginia Dept. Game and Inland Fisheries, Richmond, VA): monitored habitat use by a relict population in sub-optimal habitats. Julie Hovis (Florida Game and Fresh Water Fish Commission, Ocala, FL): inventoried a large population using global positioning units.

Nongovernmental Organizations: Roy DeLotelle (DeLotelle and Guthrie, Inc., Gainesville, FL): developed management plans for a population on private lands in central Florida. R. Todd Engstrom (Tall Timbers Research Station): monitoring a translocation experiment necessitated by a road construction project, as well as the status of a large population in south Georgia.

Management Research Programs: U.S. Forest Service: Chuck Hess (U.S. Forest Service, Crawfordville, FL and Florida State University, Tallahassee, FL): investigating food habits using a stomach and crop flushing technique. Robert G. Hooper (Southeastern Forest Experiment Station, Charleston, SC): investigating response to loss of foraging habitat in a population with high densities.

U.S. Department of Defense: Carl Petrick (Natural Resource Branch, Eglin Air Force Base, Niceville, FL) and Bob Progulske (Avon Park Air Force Base, Avon Park, FL): investigating new burning programs and developing models for ecosystem management. Stephen Parris (Environmental Division, Fort Polk Air Force Base, Fort Polk, LA): monitoring the impacts of a new multi-purpose range complex.

U.S. Fish and Wildlife Service: Ralph Costa (Red-cockaded Woodpecker Coordinator, U.S. Fish and Wildlife Service, College of Forestry and Recreation Resources, Clemson University, Clemson, SC): involved with habitat conservation planning and the review of agency management programs.

University Personnel: Dixie Reaves (Dept. Economics, Duke University, Durham, NC): performing economic valuations of red-cockaded woodpecker habitat. Richard Conner and Craig Rudolph (College of Forestry, Stephen F. Austin State University, Nacogdoches, TX): investing the influence of landscape patterns on wind and beetle damage to cavity trees as well as how birds detect red heart infections. Jeffrey Walters (Department of Zoology, North Carolina State University, Raleigh, NC) and co-workers: investigating the long-term response of populations to drilled artificial cavities.

Nongovernmental Organizations: Todd Engstrom (Tall Timber Research Station, Tallahassee, FL): investigating forest management practices on private lands in Georgia and their economic impacts and effects on woodpeckers. Robert McFarlane (McFarlane and Associates, Houston, TX): investigating the relationship between woodpecker body size and foraging territory.

Management Research Needs: Under the heading of "Population Dynamics," SSN (1990) identified the following research needs: (1) survey populations and habitat on private lands, especially near sensitive populations on public lands (<50 active cavity clusters) or private lands with potentially large populations (e.g., southern Georgia and southern Florida); (2) determine whether corridors are used to move from one area to another, or from one cluster of cavities to another (see Simberloff et al. 1992); (3) determine whether barriers of more than 8 km (5 miles) are crossed; (4) ascertain what constitutes an effective barrier to movement; (5) examine the population dynamics of small/isolated groups, including regional variation and demographic characteristics; (6) collect data to verify the basis for recommending that 400 breeding pairs would maximize long-term recovery, and determine the minimum number of populations needed for recovery (also assess viability, including environmental, demographic, and genetic characteristics); (7) examine effects of competition on reproductive success; (8) evaluate the impact of cavity competitors; (9) assess the characteristics of effective dispersal perhaps with the assistance of models; (10) monitor and analyze failures of clusters of cavities; (11) examine the survival probabilities of various populations sizes as a function of isolation, habitat, and timber management; (12) analyze survival probabilities of different stands being affected by catastrophic events -- generated from existing data; and (13) conduct a detailed survey of population size.

Under the heading of "Reproduction and Nesting," SSN (1990) identified 4 management and research topics: (1) study the effects of disturbance, such as noise and physical disturbance, and the timing, frequency, and duration of such disturbances; (2) collect and evaluate information on methods or protecting red-cockaded cavity trees from southern pine beetle; (3) assess the effectiveness of artificial cavities on actually increasing habitat; and (4) determine the feasibility and impacts of translocation to unoccupied suitable habitat.

Under the heading of "Foraging," five topics were identified (SSN 1990): (1) examine regional differences in the quality of foraging habitat; (2) develop a greater understanding of the effects of prey base on the dynamics of foraging; (3) collect more information on the impacts of hardwood removal as it affects foraging; (4) determine how foraging habitat should be configured, including age and distribution of tree species, average basal area, average diameter, stems per acre, etc.; and (5) develop a better understanding of the prey base, particularly the wood-boring ant that is found in nestling diets.

Under the heading of "Integrated Management" (SSN 1990), the following topics were identified: (1) collect more information on the impacts of converting even-aged to uneven-aged stands; (2) collect more information on management in urbanizing areas; (3) conduct more research on shortleaf and loblolly systems; (4) design and conduct research on the use of an area-wide approach to management, including an assessment of ecosystem management techniques on different forest types.

Global Protection: Many to very many (13 to >40) occurrences appropriately protected and managed

Comments: Many occurrences are on protected lands that are now managed appropriately for this species. A total of 245 occurrences are listed as protected or preserved. Although 2800 to 6000 breeding groups are supported on 98 federal or state properties, most are small, isolated, or exist in poor quality habitat; only 13 public properties support more than 275 birds (USFWS 2000). Lennartz et al. (1983) estimated that more than 3,000 active sites occurred on federal lands in the early 1980s, or about 75% of all known active sites (James 1995).

Needs: See recovery plan (USFWS 2003).

Many woodpeckers are notorious for excavating holes in such things as utility poles, fence posts and even houses, but P. borealis is very restricted in habitat, limiting any negative effects on humans. In most cases, however, the pine tree that it resides in with its clan usually dies due to the extensive excavating for nests and roosting sites. (Poole et al., 1994; Skutch, 1985)

Comments: At historic abundance levels, may be an important element in the control of pine beetle populations (McFarlane 1992).

Stewardship Overview: The primary actions needed to accomplish delisting and downlisting recovery goals are (1) application of frequent fire to both clusters and foraging habitat, (2) protection and development of large, mature pines throughout the landscape, (3) protection of existing cavities and judicious provisioning of artificial cavities, (4) provision of sufficient recruitment clusters in locations chosen to enhance the spatial arrangement of groups, and (5) restoration of sufficient habitat quality and quantity to support the large populations necessary for recovery (USFWS2003).

Management centers on maintaining old-growth pine forests and establishing an effective prescribed burning program. Minimum tree ages should be 100-125 years for longleaf pine, 80-150 years for shortleaf, and 80- 120 years for loblolly. Even older age classes are preferred. The frequency of prescribed burns should be approximately every 2-6 years in longleaf systems and 2-5 years in loblolly and shortleaf systems. Burns conducted in spring and summer are most effective in controlling hardwood encroachment. Longer burn intervals may be needed occasionally to allow seedling pines to establish.

Maintenance of existing nesting/roosting cavities is best accomplished by eliminating hardwoods in the vicinity of cavities, protecting trees against infestation by southern bark beetles (Dendroctonus frontalis), ensuring that old-growth conditions persist near cavity trees, and placing restrictor plates over cavities. Expanding the number of active cavities can be accomplished by creating artificial cavities.

Preserves large enough to support more than 25 active clusters will be stable for long periods of time and probably require infrequent intervention (so long as optimal habitat conditions are maintained). Preserves of approximately 1,000-4,000 ha (2,470-9,890 acres) have the capacity to support populations of this size. However, smaller populations (protected or not) may harbor unique or unusual genetic characteristics and also help to guard against catastrophic events. Owing to the limited dispersal capabilities evident in this species, many management areas should be considered effectively isolated.

Species Impact: Cavities constructed by red-cockaded woodpeckers weaken pine trees and make them more susceptible to wind damage (Engstrom and Evans 1990). Wind damage is a major source of natural mortality for older longleaf pines (Platt et al. 1988). Cavities may also provide an entry point for diseases (Jackson 1977), although this point is debated (Conner and Locke 1982). These impacts on pines may be offset by the fact that maintenance of habitat conditions for red-cockaded woodpeckers may reduce infestations by the southern pine beetle and the incidence of certain diseases of young pines (Berlanger et al. 1988).

Management for red-cockaded woodpeckers may benefit native plants (Platt et al. 1988a), as well as various migratory and nonmigratory birds (but compare Brennan et al., in press), including some that are known to be declining (e.g., brown-headed nuthatch and Bachman's sparrow, Aimophila aestivalis) (Robbins et al. 1987).

Comments: This species exhibits little geographic variation in morphology (Mengel and Jackson 1977). Subspecies recognized in AOU (1957) currently are regarded as not worthy of recognition.

Relative to other species of birds, there is a large among-population component of genetic variance; genetic distance among populations increases with geographic distance; small populations should not be considered "lost causes" from a genetic standpoint--they are important as reservoirs of unique genetic combinations and help facilitate gene flow among larger populations (Stangel et al. 1992).

Jackson (1971) proposed that a population of hairy woodpeckers was isolated in Florida during an inter-glacial period and eventually gave rise to the red-cockaded woodpecker. This proposition is based on morphological comparisons of isolated populations of hairy woodpeckers, which exhibit phenotypic characteristics similar to red-cockaded woodpeckers.