Setophaga pinus - (Linnaeus, 1776)
Pine Warbler
Other English Common Names: pine warbler
Synonym(s): Dendroica pinus (Linnaeus, 1776)
Taxonomic Status: Accepted
Related ITIS Name(s): Dendroica pinus (A. Wilson, 1811) (TSN 178914)
French Common Names: Paruline des pins
Spanish Common Names: Chipe Pinero
Unique Identifier: ELEMENT_GLOBAL.2.101876
Element Code: ABPBX03170
Informal Taxonomy: Animals, Vertebrates - Birds - Perching Birds
 
Kingdom Phylum Class Order Family Genus
Animalia Craniata Aves Passeriformes Parulidae Setophaga
Genus Size: D - Medium to large genus (21+ species)
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Concept Reference
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Concept Reference: American Ornithologists' Union (AOU). 1998. Check-list of North American birds. Seventh edition. American Ornithologists' Union, Washington, D.C. [as modified by subsequent supplements and corrections published in The Auk]. Also available online: http://www.aou.org/.
Concept Reference Code: B98AOU01NAUS
Name Used in Concept Reference: Dendroica pinus
Taxonomic Comments: Phylogenetic analyses of sequences of mitochondrial and nuclear DNA (Lovette et al. 2010) indicate that all species formerly placed in Dendroica, one species formerly placed in Wilsonia (citrina), and two species formerly placed in Parula (americana and pitiayumi) form a clade with the single species traditionally placed in Setophaga (ruticilla). The generic name Setophaga has priority for this clade (AOU 2011).
Conservation Status
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NatureServe Status

Global Status: G5
Global Status Last Reviewed: 09Apr2016
Global Status Last Changed: 03Dec1996
Ranking Methodology Used: Ranked by inspection
Rounded Global Status: G5 - Secure
Reasons: Large breeding range in eastern North America; common in many areas; increasing in recent decades.
Nation: United States
National Status: N5B,N5N (05Jan1997)
Nation: Canada
National Status: N5B,N5M (15Jan2018)

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Alabama (S5), Arkansas (S4), Colorado (SNA), Connecticut (S5B), Delaware (S5B,S5N), District of Columbia (S1B,S1S3N), Florida (SNR), Georgia (S5), Illinois (S3S4), Indiana (S3B), Iowa (S1N), Kentucky (S4S5B), Louisiana (S5), Maine (S5B), Maryland (S4B,S2N), Massachusetts (S4B), Michigan (S5), Minnesota (SNRB), Mississippi (S5B,S5N), Missouri (SNRB,SNRN), Nebraska (SNRN), New Hampshire (S5B), New Jersey (S4B,S4N), New York (S5B), North Carolina (S5B,S4N), Ohio (S3S4), Oklahoma (S4), Pennsylvania (S4B), Rhode Island (S4B), South Carolina (SNR), South Dakota (SNA), Tennessee (S5), Texas (S5B), Vermont (S4B), Virginia (S5), West Virginia (S2N,S4B), Wisconsin (S4B)
Canada Manitoba (S2B), New Brunswick (S5B,S5M), Nova Scotia (S1B), Ontario (S5B), Prince Edward Island (SNA), Quebec (S4B)

Other Statuses

IUCN Red List Category: LC - Least concern

NatureServe Global Conservation Status Factors

Range Extent: >2,500,000 square km (greater than 1,000,000 square miles)
Range Extent Comments: BREEDING: Subspecies PINUS from southeastern Manitoba east across Canada to southwestern New Brunswick, south to eastern Oklahoma and Texas, Gulf Coast, and southern Florida (AOU 1998). However, very local or absent within a broad section of land running east-west from western Ohio, Indiana, Illinois, and Iowa. A disjunct breeding population occurs in the "Lost Pines" area of east-central Texas (Bastrop, Caldwell, and Fayette counties). NON-BREEDING: Subspecies PINUS in the southeastern U.S. from eastern Maryland and Delaware to eastern edge of Oklahoma (Ouachita Mountains) and Texas south through the breeding range; rare to casual in northeastern U.S., Canadian provinces, southern Texas, and the Florida Keys (AOU 1998). Very few records for vagrants (mostly in winter) in Belize, Bermuda, Costa Rica, Cuba, Greenland (October), Jamaica, and northeastern Mexico. Scattered records (mostly spring and fall) in the western U.S. RESIDENT: Subspecies ACHRUSTERA in the Bahamas on Grand Bahama, Abaco, Andros, and New Providence. Subspecies CHRYSOLEUCA in the highlands of Hispaniola in western Haiti and eastern Dominican Republic (Dunn and Garrett 1997).

Number of Occurrences: 81 to >300

Population Size: 2500 to >1,000,000 individuals
Population Size Comments: No data are available, but total abundance certainly is at least several thousand individuals.

Overall Threat Impact Comments: Some forest management practices, such as clearcutting, should adversely affect the warbler because of its dependence on forest habitat. While it may be found during the breeding season in younger forests, those individuals are not necessarily breeding or breeding successfully. In Virgina, Conner et al. (1979) examined relative bird abundance in pine-oak forest clearcuts of differing ages. The species was found in increasing abundance in clearcuts greater than 10 years old and were most abundant in mature 80-year-old stands. Single-tree and group-selection cutting, while removing fewer canopy trees from forest areas, may cause increased nest predation from birds and mammals, and nest parasitism from brown-headed cowbirds (MOLOTHRUS ATER). Spread of suburban areas in pine forest regions could also cause local declines or extirpation through increased fragmentation and/or loss of forest habitat. Spraying of pesticides is of concern. Whitmore et al. (1993) found that warblers in forests treated for gypsy moths with Diflubenzuron (Dimilin [registered trademark]) had significantly lower fat reserves than birds in an untreated forest. On the same study site, Sample et al. (1993) showed no significant dietary differences of warblers between treated and untreated forests, and birds consumed the same biomass of prey on both treatments. Casualties have been recorded in areas where DDT was used to control Dutch elm disease in the 1950s (Wallace et al. 1961), and where arsenical spray had been used (Barbour 1937). Nocturnal migrants are killed at television and radio towers, though this species seems less common in kills than other species. In a 25-year study at a north Florida tower, 217 individuals were found dead (Crawford 1981). Johnston and Haines (1957) reported 106 pine wrblers killed at television towers, tall buildings, and airport ceiliometers during two nights of migration in the eastern U.S. in October 1954. Predators are not well known. Egg predators probably include blue jay (CYANOCITTA CRISTATA), American crow (CORVUS BRACHYRHYNCHOS), common grackle (QUISCALUS QUISCULA), red squirrel (TAMIASCIURUS HUDSONICUS), gray squirrel (SCIURUS CAROLINENSIS), raccoon (PROCYON LOTOR), opossum (DIDELPHIS VIRGINIANUS), and snakes. Nestling predators probably are similar, but may include domestic cats, owls, and hawks. A blue jay was observed consuming a nestling in Arkansas (Rodewald, pers. obs.). Sharp-shinned (ACCIPITER STRIATUS) and Cooper's (A. COOPERII), hawks probably prey on adults. Adults are known to mob eastern screech-owl (OTUS ASIO) (Rodewald, pers. obs.) and chuck-will's-widow (CAPRIMULGUS CAROLINENSIS) (Ficken et al. 1967), indicating those are also potential predators of adults. Reported infrequently as hosts for parasitic brown-headed cowbirds (MOLOTHRUS ATER). Friedmann and Kiff (1985) mentioned only 15 known cases of nest parasitism by cowbirds on pine warblers. Given that little is known about the breeding biology of warblers, it seems likely that it is a more frequent host of cowbirds than currently known. There is also a record of an adult female cowbird removing four warbler young from a nest and dropping them to the ground (Beane and Alford 1990). Reed (1992) used a classification system to rank neotropical migrant birds based on their relative susceptibility to extinction. Based on considerations of range, habitat specificity, distribution, and population size, the pine warbler received a "5" on a scale from 1-8, with 1 being the rating of highest concern.

Short-term Trend: Increase of >10%
Short-term Trend Comments: North American Breeding Bird Survey (BBS) data indicate a significant population increase in North America: 65% increase between 1966 and 1993, 30% increase from 1984 to 1993 (Price et al. 1995). Range-wide BBS data indicate a significant population increase of 1.6% per year during the period 1966-1994 (Sauer et al. 1997). During 1966-1979, the species showed a nonsignificant range-wide decrease of 0.8% per year (Peterjohn and Sauer 1993), and from 1980-1994 it significantly increased 2.7% per year. Witham and Hunter (1992) analyzed 1966-1987 BBS data from the northern New England region and reported a significant population increase of 5.1% per year (P < 0.05); in central New England the population increased by 8.6% per year (P < 0.01). No large-scale historical changes in distribution are known. Nesting range in Connecticut, Long Island, and locally elsewhere in the eastern United States has decreased as a result of habitat loss (Bull 1974, Zeranski and Baptist 1990). Kerlinger and Doremus (1981) reported a local extirpation in an isolated area of pine barrens near Albany, New York, even though the species was apparently not uncommon there in 1954 (Bull 1974). In Huron County, southeastern Michigan, was abundant in the early 1900s, but was not recorded in the area by the state breeding bird atlas (Brewer et al. 1991). However, in southeast Ohio, the range expanded in the 20th century in the unglaciated Allegheny Plateau (Peterjohn and Rice 1991). In areas where deciduous forest has been converted to pine plantations, such as in areas of the Ozark Highlands (e.g., Smith and Petit 1988), warblers have undoubtedly increased.

Other NatureServe Conservation Status Information

Distribution
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Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) BREEDING: Subspecies PINUS from southeastern Manitoba east across Canada to southwestern New Brunswick, south to eastern Oklahoma and Texas, Gulf Coast, and southern Florida (AOU 1998). However, very local or absent within a broad section of land running east-west from western Ohio, Indiana, Illinois, and Iowa. A disjunct breeding population occurs in the "Lost Pines" area of east-central Texas (Bastrop, Caldwell, and Fayette counties). NON-BREEDING: Subspecies PINUS in the southeastern U.S. from eastern Maryland and Delaware to eastern edge of Oklahoma (Ouachita Mountains) and Texas south through the breeding range; rare to casual in northeastern U.S., Canadian provinces, southern Texas, and the Florida Keys (AOU 1998). Very few records for vagrants (mostly in winter) in Belize, Bermuda, Costa Rica, Cuba, Greenland (October), Jamaica, and northeastern Mexico. Scattered records (mostly spring and fall) in the western U.S. RESIDENT: Subspecies ACHRUSTERA in the Bahamas on Grand Bahama, Abaco, Andros, and New Providence. Subspecies CHRYSOLEUCA in the highlands of Hispaniola in western Haiti and eastern Dominican Republic (Dunn and Garrett 1997).

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map
NOTE: The maps for birds represent the breeding status by state and province. In some jurisdictions, the subnational statuses for common species have not been assessed and the status is shown as not-assessed (SNR). In some jurisdictions, the subnational status refers to the status as a non-breeder; these errors will be corrected in future versions of these maps. A species is not shown in a jurisdiction if it is not known to breed in the jurisdiction or if it occurs only accidentally or casually in the jurisdiction. Thus, the species may occur in a jurisdiction as a seasonal non-breeding resident or as a migratory transient but this will not be indicated on these maps. See other maps on this web site that depict the Western Hemisphere ranges of these species at all seasons of the year.
Endemism: occurs (regularly, as a native taxon) in multiple nations

U.S. & Canada State/Province Distribution
United States AL, AR, CO, CT, DC, DE, FL, GA, IA, IL, IN, KY, LA, MA, MD, ME, MI, MN, MO, MS, NC, NE, NH, NJ, NY, OH, OK, PA, RI, SC, SD, TN, TX, VA, VT, WI, WV
Canada MB, NB, NS, ON, PE, QC

Range Map
Note: Range depicted for New World only. The scale of the maps may cause narrow coastal ranges or ranges on small islands not to appear. Not all vagrant or small disjunct occurrences are depicted. For migratory birds, some individuals occur outside of the passage migrant range depicted. For information on how to obtain shapefiles of species ranges see our Species Mapping pages at www.natureserve.org/conservation-tools/data-maps-tools.

Range Map Compilers: NatureServe, 2002; WILDSPACETM 2002


U.S. Distribution by County Help
State County Name (FIPS Code)
ID Fremont (16043), Latah (16057)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
17 Lower Henrys (17040203)+, Palouse (17060108)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: A 14-cm-long bird (warbler).
General Description: Slightly larger than most other wood-warblers (13-14 cm length, 9.4-15.1 g). Adult males have unstreaked, olive-green upperparts, a yellow throat and breast with indistinct black streaking on sides of the breast, a white belly and undertail coverts, dark wings with faint bluish tinge, and two broad whitish wing-bars. Adult females are duller and variable, but always with browner or grayer upperparts, a paler yellow throat and breast, duller white belly and undertail coverts. Bill is black and relatively heavy for a warbler. Immature is like adult female, but brownish gray above, breast, belly, and undertail coverts are washed with buff. Juvenile has grayish brown head and upperparts; grayish underparts are washed with buff on breast, belly, undertail coverts, and wingbars (Rodewald et al., in press).
Diagnostic Characteristics: Bay-breasted warbler (DENDROICA CASTANEA) and blackpoll warbler (D. STRIATA) in winter plumage are sometimes confused with the pine warblers. Pine warblers differ from those species in having a heavier-bodied and larger-billed appearance, unstreaked upperparts, a darker face contrasting with paler throat, a longer tail, and narrow and duller edgings on tertials. In addition, pine warblers typically show a pale yellowish or grayish area extending up the sides of the neck, which contrasts with the slightly darker face. Pine warblers have dark legs and feet (adult blackpolls usually have yellowish to pinkish legs) and are usually darker green in upperparts than are bay-breasted warblers.
Reproduction Comments: Perhaps due to the usual nesting high in trees, there are little data on reproduction. Breeding territories are established from late winter in the south to spring farther north. Breeding can begin in early-March in deep southern populations, later in the north, and may extend to early August. Based on 226 clutches from throughout the range, McNair (1987) reported a mean clutch initiation date of 20 April +/- 25 days (standard deviation). Median initiation date was 12 April; clutches were initiated between 7 March and 7 July. During an early spring in Georgia, one unfinished nest was found on 17 February (Burleigh 1958); however, most nest building does not begin until March in southern states. In more northerly states, nesting begins in April, May, or early June.

Males establish territories through persistent singing, continuous presence, and chasing or attacking intruding birds. Fights and chases among males become less common as season progresses, at which time males seem to maintain territories primarily through singing.

Clutch size is three to five (usually four). Although it is widely mentioned that the species is double-brooded, and even triple-brooded (Potter et al. 1980), there are no data to support those claims. It is almost certain the species can raise more than one brood per year, especially in southern states. Incubation, primarily by the female (male occasionally assists), lasts 12-13 days. Males feed the female on the nest. Young are fed by both parents during nestling and fledgling stages. The period of time parents feed young after fledging is unknown. Birds reach sexual maturity within one year.

Ecology Comments: Various population density figures have been reported in the literature. In Georgia, Johnston and Odum (1956) recorded 0.4 territorial males per ha in 25-year-old pine forest, 0.85 males per ha in 35-year-old forest, 1.06 males per ha in 60-year-old forest, and 1.36 males per ha in 100-year-old forest. In Texas, Dickson and Segelquist (1979) reported 0.2 territorial males per ha in pine-hardwood sapling stands, 0.55 males per ha in pine pole stands, 0.35 males per ha in pine-hardwood pole stands, 0.50 males per ha in pine saw timber stands, and 0.20 males per ha in pine-hardwood stands. In pine plantations in southern Illinois, where the species is far less common, Graber et al. (1983) reported 2.2 males per 40.5 ha. In Maryland, Stewart and Robbins (1952) recorded breeding densities of 1.9 territorial males per ha in an immature loblolly-shortleaf pine forest, 0.5 males per ha in pine-oak forest (pitch and Virginia pines, and southern red oak (QUERCUS FALCATA), and 0.25 males per ha in a mature Virginia pine forest.

Winter bird count data from the Archbold Biological Station in central Florida (December 1993), indicate a density of approximately 2.35 birds per ha (J. Fitzpatrick, in litt.). In winter, it can be abundant in pine forests of the southeast where large mixed-species flocks may contain 50-100 or more pine warblers. In fall and winter, mixed-species flocks in southeastern pine forests usually form around Carolina chickadees (POECILE CAROLINENSIS) and tufted titmice (BAEOLOPHUS BICOLOR), and often include woodpeckers, brown-headed nuthatch (SITTA PUSILLA), eastern bluebird (SIALIUS SIALIA), kinglets (REGULUS spp.), and, farther south, blue-gray gnatcatcher (POLIOPTILA CAERULEA), blue-headed vireo (VIREO SOLITARIUS), yellow-rumped (DENDROICA CORONATA) and other warblers, and chipping sparrow (SPIZELLA PASSERINA) (Gaddis 1983, Morse 1970). Warblers in fall and winter flocks can be notoriously aggressive, with males frequently fighting, chasing, or supplanting other males and females (Morse 1974). Winter mixed-species flocks in north-central Florida contained pine warblers 65% of the time and had a mean of 2.6 individuals per flock (+ or - 1.8 SE) (Gaddis 1983).

There are few data on territory size, but size probably varies considerably depending on habitat quality. In pine-oak forest in northwestern Arkansas, two pairs held territories approximately 1.0 ha in size (Rodewald, pers. obs.). Howe (1979) observed a pair in Minnesota nest building on a 0.1-ha lake island located 450 m from shore, indicating territories can be quite small in some cases. However, warblers were more regularly recorded on 1.0-ha lake islands. In oak-pine forests with low percentages of pines, may utilize only a small proportion of a much larger territory, typically moving from pine tree to pine tree and passing over deciduous trees (Morse 1974).

Interspecifically aggressive towards many bird species, especially yellow-throated warblers (D. DOMINICA) during the breeding season on Delmarva peninsula in Maryland. Pine warblers typically prevail in aggressive encounters, and were even recorded displaying towards and countersinging with yellow-throated warblers (Ficken et al. 1968, Morse 1974). Aggressive behavior towards yellow-throated warblers has also been noted in northwestern Arkansas (Rodewald, pers. obs.). Brown-headed nuthatches flocking with pine warblers in Louisiana foraged heavily on distal parts of limbs and twigs, whereas warblers foraged in areas near tree trunks. In the absence of one another, the two species exhibited similar foraging distributions, indicating that each has an influence on the foraging behavior of the other (Morse 1967).

Individuals wintering in southern forests are susceptible to extremes in weather and temperature. After 5 inches of snow and near-zero temperatures in coastal South Carolina in February 1899, Wayne (1899) reported finding countless dead birds of 16 species, including many pine warblers, a species he described as "decimated" by the cold.

Non-Migrant: Y
Locally Migrant: Y
Long Distance Migrant: Y
Mobility and Migration Comments: Migratory in the northern half of the range. In subspecies PINUS, spring migration occurs primarily from late February to mid-May. Fall migration begins in September in northern populations, with the majority of migration occurring in October. Birds in some southern populations may not migrate.
Terrestrial Habitat(s): Forest - Conifer, Forest - Mixed, Woodland - Conifer, Woodland - Mixed
Habitat Comments: Strongly associated with presence of pine and pine-hardwood forest during the breeding and winter seasons. A common breeder in most pine forests of the southeastern United States and in areas with pines in southeast Canada and the northeastern United States, but usually at lower densities, less common as a breeder in white pine forest areas. The highest numbers seem to occur where pure stands of pine are found; less abundant as the proportion of hardwood tree species increases. Birds are rarely found in deciduous forest, scrub, and thickets, except during migration and winter.

Breeding occurs in a wide variety of pine forest types but not in other conifer forests (e.g., spruce (PICEA spp.), fir (ABIES spp.), larch (LARIX spp.), or hemlock (TSUGA spp.). In the north-central and northeastern U.S. and Canadian provinces, breeding occurs in stands of red (PINUS RESINOSA), pitch (P. RIGIDA), jack (P. BANKSIANA) and white (P. STROBUS) pines (white pine also being used in the Appalachians). In southern states, breeding and winter habitat consists of stands of shortleaf (P. ECHINATA), longleaf (P. PALUSTRIS), loblolly (P. TAEDA), Virginia (P. VIRGINIANA), and slash (P. ELLIOTTII) pines. Breeding occurs less frequently in sand (P. CLAUSA) (Stevenson and Anderson 1994) and pond pines (P. SEROTINA) (Schroeder 1985) in the southeastern U.S. All forest types used may be mixed with varying proportions of hardwood species. Nesting may occur in areas of primarily deciduous forest where small groves of pines are present. Adapts well to pine plantations, which are used for breeding throughout the range. In Florida, Repenning and Labisky (1985) did not record breeding warblers in 1-, 10-, and 24-year-old slash pine plantations, but recorded 8 birds per sq km in 40-year-old plantation forests. However, they found that the species did use 1-year-old (3 birds per sq km), 24-year-old (20 birds per sq km), and 40-year-old (86 birds per sq km) pine plantations during winter.

In winter, birds commonly forage in large mixed-species flocks in southern pine forests when numbers increase because of birds migrating from farther north. At that time, flocks may forage in forest leaf litter, or in fields and pastures, usually in the vicinity of forest edge.

Density of pine warblers is inversely related to percent of deciduous vegetation within a stand (Schroeder 1985). In a breeding habitat suitability model developed by Schroeder (1985), three main habitat variables of importance were identified: percent tree canopy closure (excluding white, sand, and pond pines), successional stage of the stand, and percent of dominant canopy pines with deciduous understory in the upper one-third layer. Optimal nesting habitat was provided by pure, dense, mature pine stands (excluding pine species mentioned above) that lack a tall deciduous understory. One shortcoming of this model, however, is that warblers do use white pine forest types for nesting; they simply tend to be less common in those pine habitats.

Conner et al. (1983) reported that mature pine forests were favored in east Texas, with increasing abundance as the proportion of pole-size pines and vegetation height increased. Tree and shrub species diversity, and foliage density at different heights had little effect. In addition, stands of sapling-sized pines were avoided.

In eastern Tennessee, Anderson and Shugart (1974) found that distribution was influenced by several habitat variables, the strongest of which was related to average size of understory vegetation, number of canopy trees, and average size of canopy vegetation. In this area, birds selected areas with sparse understory and a dense canopy. In the Ouachita Mountains of Arkansas, Wilson et al. (1995) found significantly higher densities in forests with a more open midstory, lower canopy coverage, lower basal area of conifers and hardwoods, and dense ground cover of grasses, shrubs, vines, and forbs.

Nesting occurs typically in pine trees in forest, rarely in deciduous trees within pine forest. Nests usually are placed on a horizontal branch or among foliage at a branch tip, usually 8-20 m above ground. Nests are usually well hidden and difficult to observe from the ground.

Adult Food Habits: Frugivore, Granivore, Invertivore
Immature Food Habits: Frugivore, Granivore, Invertivore
Food Comments: Arthropods are of primary importance. Lepidoptera larvae and pupae probably are the most frequent food items; other foods include Blattidae, Coccididae, Diptera, Homoptera, Hemiptera, Orthoptera, Araneae, Coleoptera, Hymenoptera, and Arachnida (Nesbitt and Hetrick 1976, Sample et al. 1993). During late summer, fall, and winter, the diet also may include fruits of bayberry (MYRICA spp.), dogwood (CORNUS FLORIDA), grape (VITUS spp.), persimmon (DIOSPYROS VIRGINIANA), sumac (RHUS spp.), and Virginia creeper (PARTHENOCISSUS QUINQUEFOLIA) (Bent 1953). This species apparently is unique among warblers in its ability to consume seeds to a large extent. Most seeds eaten are those of pines, such as longleaf, pitch, shortleaf, and loblolly. However, little information exists on the frequency of seed-eating (see Morse 1967).

Foraging consists primarily of gleaning pine foliage and bark substrates, and sometimes deciduous foliage, especially during migration. Foraging occurs commonly on leaf litter of the forest floor in fall and winter, and sometimes in open habitats near forest edge. Sometimes flycatching is used to capture aerial prey items. Resident birds on Grand Bahama island foraged heavily on pine foliage and infrequently on bark surfaces, whereas birds used pine foliage and bark substrates approximately equally on Andros island (Emlen 1981).

Foraging movements are relatively slowly for a wood warbler, consisting of hopping along tree limbs and branches and searching bark and foliage substrates while ducking twigs and foliage. This warbler regularly searches bark of tree trunk by foraging at bases of branches or landing on trunk. This behavior also has been recorded during spring migration in deciduous forest before leaves emerge. Frequently, it forages in pine foliage at branch tips, moving from one branch tip to the next, sometimes hanging sideways or upsidedown to procure prey items.

Birds in the New Jersey Pine Barrens may respond to seasonal changes in arthropod abundance in oaks (Brush and Stiles 1990). In May, similar densities of pine warblers were found in two oak-pine forests, one of which was dominated by pines. However, when arthropod abundances were higher in the pine-dominated forest in June and July, significantly higher densities of birds were recorded in that forest. In pine forests during the breeding season, birds tended to forage higher, often at tips of branches in the canopy. Additional information on foraging can be found in Rodewald et al. (in prep.).

Adult Phenology: Diurnal
Immature Phenology: Diurnal
Phenology Comments: No quantitative information is available on diurnal fluctuations in activity, but birds seem more active from early to mid-morning, and late afternoon to early evening.
Length: 14 centimeters
Weight: 12 grams
Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: BBS data indicate that populations were increasing during the years of 1966-1991 (Peterjohn and Sauer 1993). Thus, this is not currently a species of high management concern in much of its range. However, in areas where native pine forests have been severely reduced in extent or marginalized for breeding by fragmentation, habitat management for this species may be necessary to maintain viable populations. Breeding habitat can be managed through controlled burning and/or forest midstory thinning.
Restoration Potential: With populations apparently increasing, the potential of restoring this species to higher population levels is quite good. Some habitat will be lost in the future through expanding urban/suburban areas. However, changes in land and forest management practices could expand available habitat and improve existing habitats to make them more productive for this and possibly other species (e.g., Bachman's sparrow (AIMOPHILA AESTIVALIS), brown-headed nuthatch, and red-cockaded woodpecker (PICOIDES BOREALIS) in southern pine forests).
Preserve Selection & Design Considerations: Given the present knowledge of the natural history of the warbler, it is difficult to make entirely informed suggestions about characteristics of preserve design. There is strong indication that large forest area is important to sustain viable populations. Whitcomb et al. (1981) considered it a forest-interior species. Schroeder (1985) mentions a minimum estimate of 10-15 ha of forest habitat as required to sustain a breeding population and Robbins (1979) estimated the minimum area to be 30 ha. However, Robbins et al. (1989) found no significant relationship between probability of occurrence and forest area for pine warblers. While this study found no indications that forest isolation is of importance to this species, it is likely that high isolation of fragments could lead to declines or extirpation in isolated patches. Lynch and Whigham (1984) showed that even a modest degree of habitat isolation reduced abundance of some migratory bird species in Maryland.

In a old-growth hammock in Florida, Noss (1991) recorded significantly higher numbers of warblers closer to edges than in the forest interior during the breeding season, and considered it an "edge-attracted species." While this species may be attracted to edge habitats in some areas, edge habitats of small forest blocks probably do not support viable populations. It is advisable that pine forest preserves be subjected to as little internal fragmentation as possible; edges may cause increases in nest predation and parasitism.

Management Requirements: In southern forests, controlled burning that creates habitat for red-cockaded woodpecker, Bachman's sparrow, and brown-headed nuthatch also provides habitat for warblers. Wilson et al. (1995) documented higher densities of the species on stands subjected to wildlife stand improvement in Arkansas (includes prescribed fire, thinning of midstory and codominant trees). In pine forests where fire suppression is practiced, expanding hardwood midstory should decrease warbler abundance, and may, in some areas, exclude the species. Manual hardwood midstory cutting (without burning) would also likely create adequate habitat for the species.
Monitoring Requirements: Pine warblers and other bird species can be surveyed during the breeding season from April-June (optimal sampling period will depend on region) using fixed-radius point-count (Hutto et al. 1986) or strip transect (Emlen 1977) methods. Counts should be conducted in morning hours from dawn until approximately 10:00 hours.
Management Research Needs: 1) Pine warblers occupy a wide variety of pine forest types throughout their range. Those forests vary somewhat in vegetative composition and the dominant pine species present. While habitat-relationships have been studied in some areas of the range, a range-wide survey of habitats and habitat preferences would be useful from a management perspective.

2) Surprisingly little is known about basic natural history. Information on nesting biology (including nest-site selection) and territory/home range size would be particularly useful.

Biological Research Needs: 1) During winter in pine forests of the southeastern U.S., when many warblers are found in mixed-species flocks, a significant number of individuals, primarily males, are regularly observed outside flocks. Are these birds nonmigratory residents maintaining winter territories? Interestingly, both solitary males and males in flocks sing during winter. Studies with color-banded populations could yield interesting information on flocking behavior and possible winter territoriality.

2) Seed-eating by this species is of much interest. The pine warbler is apparently the only warbler that consumes seeds (primarily pine seeds) in any significant amount. However, few data exist on how regular this behavior may be or on the importance of seeds in the fall/winter diet. In addition, little is known about how this primarily insectivorous bird may make the seasonal physiological changes required to allow for the digestion of seeds.

Population/Occurrence Delineation
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Group Name: Passerines

Use Class: Breeding
Subtype(s): Foraging Area, Nest Site, Nesting Colony
Minimum Criteria for an Occurrence: Evidence of historical breeding, or current and likely recurring breeding, at a given location, minimally a reliable observation of one or more breeding pairs in appropriate habitat. Be cautious about creating EOs for observations that may represent single breeding events outside the normal breeding distribution.

Mapping Guidance: Breeding occurrences include nesting areas as well as foraging areas.

For swallows and other species that have separate nesting and foraging areas, separations are based on nest sites or nesting areas, not to locations of foraging individuals. For example, nesting areas separated by a gap larger than the separation distance are different occurrences, regardless of the foraging locations of individuals from those nesting areas. This separation procedure is appropriate because nesting areas are the critical aspect of swallow breeding occurrences, tend to be relatively stable or at least somwhat predictable in general location, and so are the basis for effective conservation; foraging areas are much more flexible and not necessarily static.

Separation Distance for Unsuitable Habitat: 5 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Significant dispersal and associated high potential for gene flow among populations of birds separated by tens of kilometers (e.g., Moore and Dolbeer 1989), and increasing evidence that individuals leave their usual home range to engage in extrapair copulations, as well as long foraging excursions of some species, make it difficult to circumscribe occurrences on the basis of meaningful population units without occurrences becoming too large. Hence, a moderate, standardized separation distance has been adopted for songbirds and flycatchers; it should yield occurrences that are not too spatially expansive while also accounting for the likelihood of gene flow among populations within a few kilometers of each other.

Be careful not to separate a population's nesting areas and foraging areas as different occurrences; include them in the same occurrence even if they are more than 5 km apart. Mean foraging radius (from nesting area) of Brown-headed Cowbird females was 4.0 kilometers in California, 1.2 kilometers in Illinois-Missouri (Thompson 1994). Yellow-headed Blackbirds, Brewer's Blackbirds, and probably Red-winged Blackbirds all forage up to 1.6 kilometers away from breeding colony (Willson 1966, Horn 1968). In one study, Brewer's Blackbirds were found as far as 10 kilometers from nesting area (Williams 1952), but this may be unusual.

For swallows and other parrerines with similar behavioral ecology, separation distance pertains to nest sites or nesting colonies, not to locations of foraging individuals. For example, nesting areas separated by a gap of more than 5 km are different occurrences, regardless of the foraging locations of individuals from those nesting areas. This separation procedure is appropriate because nesting areas are the critical aspect of swallow breeding occurrences, tend to be relatively stable or at least somwhat predictable in general location, and so are the basis for effective conservation; foraging areas are much more flexible and not necessarily static.

Be cautious about creating EOs for observations that may represent single breeding events outside the normal breeding distribution.

Unsuitable habitat: Habitat not normally used for breeding/feeding by a particular species. For example, unsuitable habitat for grassland and shrubland birds includes forest/woodland, urban/suburban, and aquatic habitats. Most habitats would be suitable for birds with versatile foraging habits (e.g., most corvids).

Date: 10Sep2004
Author: Hammerson, G.

Use Class: Migratory stopover
Subtype(s): Foraging Area, Roost Site
Minimum Criteria for an Occurrence: For most passerines: Evidence of recurring presence of migrating individuals (including historical) and potential recurring presence at a given location; minimally a reliable observation of 25 birds in appropriate habitat.

For swallows: Evidence of recurring presence of migrating flocks (including historical) and potential recurring presence at a given location; minimally a reliable observation of 100 birds in appropriate habitat (e.g., traditional roost sites).

Occurrences should be locations where the species is resident for some time during the appropriate season; it is preferable to have observations documenting presence over at least 7 days annually.

EOs should not be described for species that are nomadic during nonbreeding season: e.g., Lark Bunting.

Be cautious about creating EOs for observations that may represent single events.

Separation Distance for Unsuitable Habitat: 5 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Separation distance somewhat arbitrary but intended to define occurrences of managable size for conservation purposes. Occurrences defined primarily on the basis of areas supporting concentrations of birds, rather than on the basis of distinct populations.

For swallows and other species with similar behavioral ecology, the separation distance pertains to communal roost sites rather than to foraging areas; the former tend to be more stable and specific over time than the latter.

Date: 03Sep2004
Author: Hammerson, G., and S. Cannings

Use Class: Nonbreeding
Subtype(s): Foraging Area, Roost Site
Minimum Criteria for an Occurrence: Any area used traditionally in the nonbreeding season (used for populations that are not resident in a location year-round). Minimally, reliable observations of 10 or more individuals in appropriate habitat for 20 or more days at a time. For G1-G3 species, observations of fewer individuals could constitute an occurrence of conservation value. Sites used during migration should be documented under the 'migratory stopover' location use class.

Separation Distance for Unsuitable Habitat: 5 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Separation distance is necessarily arbitrary but attempts to balance the high mobility of birds with the need for occurrences of reasonable spatial scope. Note that a population's roost sites and foraging areas are parts of the same occurrence, even if they are more than 5 km apart.

For swallows and other species with similar behavioral ecology, the separation distance pertains to communal roost sites rather than to foraging areas; the former tend to be more stable and specific over time than the latter.

Date: 03Sep2004
Author: Hammerson, G.

Use Class: Nonmigratory
Minimum Criteria for an Occurrence: Occurrences are based on evidence of historical presence, or current and likely recurring presence, at a particular location. Such evidence minimally includes collection or reliable observation and documentation of one or more individuals in or near appropriate habitat.

These occurrence specifications are used for nonmigratory populations of passerine birds.

Separation Barriers: None.
Separation Distance for Unsuitable Habitat: 5 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Significant dispersal and associated high potential for gene flow among populations of birds separated by tens of kilometers (e.g., Moore and Dolbeer 1989), and increasing evidence that individuals leave their usual home range to engage in extrapair copulations, as well as long foraging excursions of some species, make it difficult to circumscribe occurrences on the basis of meaningful population units without occurrences becoming too large. Hence, a moderate, standardized separation distance has been adopted for songbirds and flycatchers; it should yield occurrences that are not too spatially expansive while also accounting for the likelihood of gene flow among populations within a few kilometers of each other.

Be careful not to separate a population's nesting areas and breeding-season foraging areas as different occurrences; include them in the same occurrence even if they are more than 5 km apart. Blue jays have small summer home ranges but fly up to 4 kilometers to harvest mast (Tarvin and Woolfenden 1999). Flocks of pinyon jays range over 21-29 square kilometers (Ligon 1971, Balda and Bateman 1971); nesting and foraging areas may be widely separated. Tricolored blackbirds forage in flocks that range widely to more than 15 kilometers from the nesting colony (Beedy and Hamilton 1999).

Unsuitable habitat: Habitat not normally used for breeding/feeding by a particular species. For example, unsuitable habitat for grassland and shrubland birds includes forest/woodland, urban/suburban, and aquatic habitats. Most habitats would be suitable for birds with versatile foraging habits (e.g., most corvids).

Date: 10Sep2004
Author: Hammerson, G.
Notes: These specs pertain to nonmigratory species.
Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 16Feb1996
NatureServe Conservation Status Factors Author: Dirrigl, F., Jr., P. G. Rodewald, K. G. Smith, & G. Hammerson
Management Information Edition Date: 01Oct1995
Management Information Edition Author: RODEWALD, P.G., AND K.G. SMITH; REVISIONS BY G. HAMMERSON, F. DIRRIGL, JR., AND D.W. MEHLMAN
Element Ecology & Life History Edition Date: 16Feb1996
Element Ecology & Life History Author(s): RODEWALD, P. G., K. G. SMITH, AND G. HAMMERSON

Zoological data developed by NatureServe and its network of natural heritage programs (see Local Programs) and other contributors and cooperators (see Sources).

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"Data provided by NatureServe in collaboration with Robert Ridgely, James Zook, The Nature Conservancy - Migratory Bird Program, Conservation International - CABS, World Wildlife Fund - US, and Environment Canada - WILDSPACE."

Citation for Mammal Range Maps of North America:
Patterson, B.D., G. Ceballos, W. Sechrest, M.F. Tognelli, T. Brooks, L. Luna, P. Ortega, I. Salazar, and B.E. Young. 2003. Digital Distribution Maps of the Mammals of the Western Hemisphere, version 1.0. NatureServe, Arlington, Virginia, USA.

Acknowledgement Statement for Mammal Range Maps of North America:
"Data provided by NatureServe in collaboration with Bruce Patterson, Wes Sechrest, Marcelo Tognelli, Gerardo Ceballos, The Nature Conservancy-Migratory Bird Program, Conservation International-CABS, World Wildlife Fund-US, and Environment Canada-WILDSPACE."

Citation for Amphibian Range Maps of the Western Hemisphere:
IUCN, Conservation International, and NatureServe. 2004. Global Amphibian Assessment. IUCN, Conservation International, and NatureServe, Washington, DC and Arlington, Virginia, USA.

Acknowledgement Statement for Amphibian Range Maps of the Western Hemisphere:
"Data developed as part of the Global Amphibian Assessment and provided by IUCN-World Conservation Union, Conservation International and NatureServe."

NOTE: Full metadata for the Bird Range Maps of North America is available at:
http://www.natureserve.org/library/birdDistributionmapsmetadatav1.pdf.

Full metadata for the Mammal Range Maps of North America is available at:
http://www.natureserve.org/library/mammalsDistributionmetadatav1.pdf.

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