Protonotaria citrea - (Boddaert, 1783)
Prothonotary Warbler
Other English Common Names: prothonotary warbler
Taxonomic Status: Accepted
Related ITIS Name(s): Protonotaria citrea (Boddaert, 1783) (TSN 178846)
French Common Names: paruline orangée
Spanish Common Names: Chipe Dorado
Unique Identifier: ELEMENT_GLOBAL.2.100372
Element Code: ABPBX07010
Informal Taxonomy: Animals, Vertebrates - Birds - Perching Birds
 
Kingdom Phylum Class Order Family Genus
Animalia Craniata Aves Passeriformes Parulidae Protonotaria
Genus Size: A - Monotypic genus
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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: Protonotaria citrea
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 much of eastern U.S. and adjacent southern Ontario; numerous occurrences; fairly stable population but with some indications of a small decline.
Nation: United States
National Status: N5B (19Mar1997)
Nation: Canada
National Status: N1B,NNRM (03Jan2018)

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 (S5B), Arizona (S1M), Arkansas (S4B), Colorado (SNA), Connecticut (SNA), Delaware (S4B), District of Columbia (S1B), Florida (S4B), Georgia (S4), Illinois (S5), Indiana (S4B), Iowa (S3B,S3N), Kansas (S3B), Kentucky (S5B), Louisiana (S5B), Maryland (S4B), Massachusetts (S1B,S2N), Michigan (S3), Minnesota (SNRB), Mississippi (S5B), Missouri (S4), Nebraska (S2), New Jersey (S4B), New Mexico (S4N), New York (S2B), North Carolina (S5B), Ohio (S3), Oklahoma (S4B), Pennsylvania (S2S3B), Rhode Island (S1B,S1N), South Carolina (S3B), South Dakota (SNA), Tennessee (S4), Texas (S3B), Virginia (S4), West Virginia (S2B), Wisconsin (S2B)
Canada Ontario (S1B)

Other Statuses

Canadian Species at Risk Act (SARA) Schedule 1/Annexe 1 Status: E (05Jun2003)
Committee on the Status of Endangered Wildlife in Canada (COSEWIC): Endangered (27Nov2016)
Comments on COSEWIC: Reason for designation: In Canada, this species breeds only in deciduous swamp forests in southwestern Ontario. It has shown an 80% decrease in abundance over the last 10 years and its current population is between 28 and 34 mature individuals only. Threats include loss and degradation of breeding habitat, loss of coastal mangrove forests in Central and South America where the species winters, and disturbances of habitat that result in increased nest site competition with House Wrens and increased nest parasitism by Brown-headed Cowbirds.

Status history: Designated Special Concern in April 1984. Status re-examined and designated Endangered in April 1996. Status re-examined and confirmed in May 2000, April 2007, and November 2016.

IUCN Red List Category: LC - Least concern

NatureServe Global Conservation Status Factors

Range Extent: 20,000-2,500,000 square km (about 8000-1,000,000 square miles)
Range Extent Comments: BREEDING: eastern Minnesota and Wisconsin across southern Great Lakes region to northern New Jersey, south to Texas, Gulf Coast, and central Florida, west to Oklahoma and Kansas (AOU 1983). BBS data (1966-1991) indicate relatively high abundance in Louisiana, North Carolina, and Mississippi. Breeding range has expanded northward, especially in the Mississippi Valley and vicinity, and nesting recently (1992) was recorded in Rhode Island (Wadman et al., unpubl.). NON-BREEDING: Yucatan peninsula (rarely) south on Caribbean slope of Central America to Nicaragua; both slopes of Costa Rica and Panama; from Colombia east to northern Venezuela; and from the Netherlands Antilles east to Trinidad and Tobago (AOU 1983). Most common in Panama and western Colombia and northern Venezuela; barely reaches Suriname and northern Ecuador. Migrates through and apparently occasionally overwinters in West Indies. Uncommon in Puerto Rico, apparently rare in Virgin Islands (Raffaele 1983, 1989). Recorded also on the Galapagos Islands nearly 1000 km west of mainland South America (Petit and Tarvin 1990). Birdlife International (2014) estimates a range size of 1.4 million square kilometers

Area of Occupancy: >12,500 4-km2 grid cells
Area of Occupancy Comments: With an average of about 4 hectares per breeding pair, the area of occupancy should easilyi exceed 20,000 square kilometers given the estimated population size for this species.

Number of Occurrences: 81 to >300
Number of Occurrences Comments: This species has an extremely large range and populatioin size so it should have at least 81 EOs.

Population Size: >1,000,000 individuals
Population Size Comments: Partners in Flight (2013) estimates 1.6 million individuals of this species.

Number of Occurrences with Good Viability/Integrity: Many to very many (41 to >125)
Viability/Integrity Comments: Estimate based on population size and range

Overall Threat Impact Comments: The primary threat in most areas is loss of suitable habitat (Leberman 1992; Busby and Mabey, pers. comms.). Loss of old growth forest associated with riparian habitats is detrimental because older trees are more likely to develop nesting cavities. Widespread drainage of required wetland habitat is a significant threat. Significant mortality factors appear to be competition with, and nest destruction, by house wrens (TROGLODYTES AEDON) (Walkinshaw 1941, 1953, Petit 1989, Brush 1991) in northern parts of the range, brood parasitism by brown-headed cowbirds (MOLOTHRUS ATER) (Walkinshaw 1941, Bent 1953, Petit 1989, 1991), predatory mammals, flooding and predatory birds (Walkinshaw 1941), and snakes (Bent 1953, Petit 1989). Nesting success was seriously impacted by competing house wrens in eastern Iowa (Brush 1991). Where nest-boxes were available, house wrens used 90-94% of them, making them unavailable to warblers. In areas that lacked nest-boxes but had higher water levels, wrens took over fewer warbler nests, and predation on warbler nests was rare. Guillory (1987) provided evidence of competition for cavities and predation on prothonotary warblers by PEROMYSCUS spp. (white-footed and cotton mice) in Louisiana; mice destroyed nests, killed adults and nestlings, and cached food in cavities and nest boxes. Predator-free habitat with high, stable water levels and low wren populations, appears to be more beneficial than are nest-boxes placed in low-quality habitat. By placing the nest over water, may be better protected from mammalian predators (Petit 1989). As a cavity nester, is probably less vulnerable than open-nesting species to increased predation or brood parasitism caused by forest fragmentation. However, cowbird parasitism has been documented to reduce nest success by approximately 25%, primarily through egg removal by female cowbirds and lowered hatching success of warbler eggs (Petit 1991). Petit (1991) found 21% of 172 nests (in nest-boxes) parasitized by cowbirds (natural cavities were parasitized at the same rate)--a relatively high frequency of parasitism for a hole-nesting species. The impact of parasitism appeared relatively low, however, compared with other host species. Females accepted parasitism in 81% of cases and were often able to raise most of their young along with the cowbird. Petit (1991) suggested that acceptance of parasitism may be more likely to occur when females have limited opportunities to renest in a different site. Prothonotary warblers therefore may be more strongly impacted by forest management practices that alter habitat or remove valuable cavity trees; with fewer nest-sites, cowbird parasitism will have greater effects. Two females were found dead on their nests by and were believed to have died as a result of prior illness and the energetic demands of reproduction (Petit and Petit 1987).

Short-term Trend: Relatively Stable (<=10% change)
Short-term Trend Comments: North American Breeding Bird Survey (BBS) data indicate a relatively stable population, 1966-1989 (nonsignificant decline of 0.1% per year) (Droege and Sauer 1990); nonsignificant decline of 18% from 1966 to 1993, nonsignificant decline of 12% from 1984 to 1993 (Price et al. 1995). Using rangewide BBS data, Sauer and Droege (1992) found a population increase of 0.95% per year over the long-term (1966-1988) and 0.01% per year over the short-term (1978-1988); neither of these changes are statistically significant. When BBS data from only the eastern region were analyzed, the population was found to significantly increase by 1.75% per year over the long-term and by 1.06% per year (not significant) over the short-term. Data on population trends are somewhat variable depending on the period and region of data analyzed. The center of abundance is the South Atlantic Coastal Plain physiographic area, where BBS routes achieve the highest rates of detection. In this region, there are signs of decline and these signs of decline are widespread regardless of detection rates. BBS data in general show significant increases over both the last 10-year and 20-year periods (Hunter, unpubl. data). During the decade 1982-1991, BBS data show significant population increases in three states (Indiana, Mississippi, and North Carolina), significant population decreases in five states (Alabama, Arkansas, Georgia, Louisiana, and Virginia), and nonsignificant trends in the remaining 12 states where data were collected (Delaware, Florida, Illinois, Kentucky, Maryland, Michigan, Missouri, New Jersey, Oklahoma, South Carolina, Tennessee, and Texas). Other statistically significant changes are a general decline in the east, declines in two physiographic regions (Upper Coastal Plain and Mississippi Alluvial Plain), and increases in two physiographic regions (Lower Coastal Plain and Lexington Plain). James et al. (1992) analyzed BBS data for the 22-year period 1966-1987 for the southeastern and south-central U.S. and distinguished between upland and lowland forests. While populations showed no trend in upland forests, they significantly increased in lowland forests. A significant increase occurred in the Lower Coastal Plain, but a decrease occurred in the Ozark Ouachita Plateau; a major peak in numbers in the late 1970s in both the Mississippi Alluvial Plain and the Upper Coastal Plain dominates the overall trend for the region. Numbers increased in the Mississippi Alluvial Plain and the Lower Coastal Plain in the first half of the period and declined in the second half, a pattern not seen elsewhere in the region for this species. Overall, although higher at the end of the period (1987) than at the beginning (1966), numbers have been declining since the late 1970s. According to Robbins et al. (1986), a strong increase in the southeast produced a significant rise in the continental population between 1965 and 1979 despite a high degree of irregularity during the last years of this period. Because habitat for this species is restricted to wooded wetland areas, the total numbers recorded in any state were small and no significant state trends were detected. This species is one of few to reach its greatest density in the Mississippi Alluvial Plain; the largest numbers were recorded in Louisiana. Very small population (probably not much more than about 50 pairs) and limited distribution in Canada (southern Ontario), but numbers stable (McCracken 1984). Although the evidence is weak, winter populations may be declining in Puerto Rico. Faaborg and Arendt (1992) caught only three individuals between 1981 and 1990 in annual mist-netting studies in the Guanica Forest. In the three study-years previous to this period (1976, 1978, and 1980), four birds were caught.

Long-term Trend: Decline of 30-50%
Long-term Trend Comments: The latest BBS data for 1996 - 2012 shows an estimated 1.08% annual decline over the entire time period, which translates to a 40% decline (Sauer, et. al. 2014).

Intrinsic Vulnerability: Moderately vulnerable
Intrinsic Vulnerability Comments: Prothonotary Warblers are vulnerable because of habitat loss on both their breeding and wintering grounds. An estimated 90% of the orgiinal breeding habitat has been lost due to logging or farming (National Audubon Society, 2014). This species lives in mangrove forest on its wintering grounds but mangroves are being lost at the rate of 1.7% per year along the Caribbean mainland (National Audubon Society, 2014).

Environmental Specificity: Very narrow to narrow.
Environmental Specificity Comments: Breeding is highliy localized because of extreme habitat specificity: foreted areas with standing waters (Petit, 1999). Winter populations are specific to mangroves, an increasingly rare habitat.

Other NatureServe Conservation Status Information

Inventory Needs: A better inventory and study of breeding success comparing natural cavity nesting versus nest boxes would be useful (Petit, 1999).

Protection Needs: Local, volunteer-based nest-box programs for the species are becoming more common in regional and county parks to repopulate areas where populations have dwindled or disappeared (Cartwright 1997, LJP). Use of nest boxes in forested wetlands where natural nest sites have been lost over time can be important tool for enhancing declining local populations (Petitk 1999). General efforts to conserve their habitat will also benefit this species.

Distribution
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Global Range: (20,000-2,500,000 square km (about 8000-1,000,000 square miles)) BREEDING: eastern Minnesota and Wisconsin across southern Great Lakes region to northern New Jersey, south to Texas, Gulf Coast, and central Florida, west to Oklahoma and Kansas (AOU 1983). BBS data (1966-1991) indicate relatively high abundance in Louisiana, North Carolina, and Mississippi. Breeding range has expanded northward, especially in the Mississippi Valley and vicinity, and nesting recently (1992) was recorded in Rhode Island (Wadman et al., unpubl.). NON-BREEDING: Yucatan peninsula (rarely) south on Caribbean slope of Central America to Nicaragua; both slopes of Costa Rica and Panama; from Colombia east to northern Venezuela; and from the Netherlands Antilles east to Trinidad and Tobago (AOU 1983). Most common in Panama and western Colombia and northern Venezuela; barely reaches Suriname and northern Ecuador. Migrates through and apparently occasionally overwinters in West Indies. Uncommon in Puerto Rico, apparently rare in Virgin Islands (Raffaele 1983, 1989). Recorded also on the Galapagos Islands nearly 1000 km west of mainland South America (Petit and Tarvin 1990). Birdlife International (2014) estimates a range size of 1.4 million square kilometers

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, AZ, CO, CT, DC, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, MI, MN, MO, MS, NC, NE, NJ, NM, NY, OH, OK, PA, RI, SC, SD, TN, TX, VA, WI, WV
Canada ON

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


U.S. Distribution by County Help
State County Name (FIPS Code)
MI Allegan (26005), Berrien (26021), Branch (26023), Calhoun (26025), Clinton (26037), Eaton (26045), Gratiot (26057), Ingham (26065), Ionia (26067), Kalamazoo (26077), Macomb (26099), Monroe (26115), Muskegon (26121), Oceana (26127), Saginaw (26145), St. Joseph (26149), Van Buren (26159), Wayne (26163)
NE Sarpy (31153)
NJ Salem (34033)
NY Genesee (36037), Niagara (36063), Oneida (36065), Orange (36071), Orleans (36073), Oswego (36075), Suffolk (36103)
OH Delaware (39041), Franklin (39049), Gallia (39053), Geauga (39055), Hamilton (39061), Hancock (39063), Holmes (39075), Lake (39085)*, Licking (39089), Logan (39091), Lucas (39095), Ottawa (39123), Seneca (39147), Summit (39153), Wayne (39169)
PA Adams (42001)*, Beaver (42007), Berks (42011)*, Bucks (42017)*, Crawford (42039), Erie (42049), Lancaster (42071), Montgomery (42091), Potter (42105), Union (42119)*, York (42133)
RI Washington (44009)
WI Buffalo (55011), Burnett (55013), Columbia (55021), Crawford (55023), Grant (55043), Green (55045), Green Lake (55047), Iowa (55049), Jackson (55053), Jefferson (55055), La Crosse (55063), Marathon (55073), Outagamie (55087), Pepin (55091), Pierce (55093), Polk (55095), Richland (55103), Rock (55105), Sauk (55111), Shawano (55115), St. Croix (55109), Trempealeau (55121), Vernon (55123), Waupaca (55135), Wood (55141)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
01 Pawcatuck-Wood (01090005)+
02 Mohawk (02020004)+, Rondout (02020007)+, Northern Long Island (02030201)+, Southern Long Island (02030202)+, Middle Delaware-Musconetcong (02040105)+*, Schuylkill (02040203)+, Cohansey-Maurice (02040206)+, Middle West Branch Susquehanna (02050203)+, Pine (02050205)+, Lower West Branch Susquehanna (02050206)+*, Lower Susquehanna (02050306)+, Monocacy (02070009)+*
04 Upper Fox (04030201)+, Wolf (04030202)+, Little Calumet-Galien (04040001)+, St. Joseph (04050001)+, Kalamazoo (04050003)+, Upper Grand (04050004)+, Maple (04050005)+, Lower Grand (04050006)+, Pere Marquette-White (04060101)+, Muskegon (04060102)+, Shiawassee (04080203)+, Clinton (04090003)+, Huron (04090005)+, Blanchard (04100008)+, Lower Maumee (04100009)+*, Cedar-Portage (04100010)+, Sandusky (04100011)+, Cuyahoga (04110002)+, Ashtabula-Chagrin (04110003)+*, Chautauqua-Conneaut (04120101)+, Niagara (04120104)+, Oak Orchard-Twelvemile (04130001)+, Oneida (04140202)+
05 French (05010004)+, Upper Ohio (05030101)+, Shenango (05030102)+, Walhonding (05040003)+, Licking (05040006)+, Upper Scioto (05060001)+, Upper Great Miami (05080001)+, Lower Great Miami (05080002)+*, Raccoon-Symmes (05090101)+, Little Miami (05090202)+*, Middle Ohio-Laughery (05090203)+
07 Upper St. Croix (07030001)+, Lower St. Croix (07030005)+, Rush-Vermillion (07040001)+, Buffalo-Whitewater (07040003)+, La Crosse-Pine (07040006)+, Black (07040007)+, Lower Chippewa (07050005)+, Coon-Yellow (07060001)+, Grant-Little Maquoketa (07060003)+, Lake Dubay (07070002)+, Castle Rock (07070003)+, Baraboo (07070004)+, Lower Wisconsin (07070005)+, Kickapoo (07070006)+, Crawfish (07090002)+, Sugar (07090004)+
10 Big Papillion-Mosquito (10230006)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: A 14-cm bird (warbler).
General Description: Length 14 cm. Relatively large, plump, short-tailed, and long-billed compared to other Parulidae. Eyes are large, dark, and prominent. Male's head and underparts are golden yellow, fading to white undertail coverts; wings are blue-gray, without wing bars; blue-gray tail has large white patches. Female is duller, head less golden. See Kowalski (1986) for information on weights, measurements, and methods of aging.

VOCALIZATIONS: Song is a series of loud, ringing "zweet" notes (NGS 1987).

NEST: cup-shaped hollow of mosses, rootlets, twigs, and leaves, smoothly lined with fine grasses, leaf stems, and feathers; inside diameter 5.1 cm, depth 3.8 cm.

EGGS: average 18.47 x 14.55 mm; oval to short-oval; shell is smooth, somewhat glossy, creamy, boldly and liberally spotted and blotched with brown.

Diagnostic Characteristics: Distinguished from yellow warbler (DENDROICA PETECHIA) by blue-gray wings and uppertail feathers; distinguished from blue-winged warbler (VERMIVORA PINUS) by lack of a black eye line.
Reproduction Comments: In the mid-Atlantic region, nesting occurs from late April to late June, with a peak from mid-May to mid-June (see Bushman and Therres 1988). Petit (1989) found the mean initiation dates for late nests to be 7 June in 1985, and 13 June in 1987. Clutch size is 3-8 (usually 4-6) (Walkinshaw 1941, 1953; Petit 1989; Blem and Blem 1992). Incubation, by female, typically lasts 12-14 days. Young are tended by both parents, leave nest at 10-11 days. Typically, individual females produce two broods per year, sometimes one, especially in the north. Petit (1989) found most females to lay two clutches, but Blem and Blem (1992) never found the same female incubating two sets of eggs in any single year.

Reproductive success in natural cavities (e.g., Walkinshaw 1941) appears to be significantly lower than in nest-boxes (e.g., Petit 1989). Walkinshaw (1941) reported a hatching success of 38% in Michigan and 61% in Tennessee; Petit (1989) reported a hatching success of 84% and a low nestling mortality of 11%, compared to Walkinshaw's (1941) 33% in Michigan. The difference in nest success appears to be most attributable to regional differences in competition from house wrens (TROGLODYTES AEDON). Life history accounts were provided by Brewster (1878), Loucks (1894), Meyer and Nevius (1943), Bent (1953), Walkinshaw (1953), and Petit (1989).

Ecology Comments: POPULATION DENSITY: Published information on bird densities from breeding bird censuses in the southeastern U.S. between 1947 and 1979 were summarized by Hamel et al. (1982): mean (se) density was listed as 14.9 (3.9) pairs per 40 ha, with a density range of 6-28 pairs per 40 ha. Accurate measures of population densities from censusing techniques are rare for this species because few studies of this nature occur in swamp forests. Encouragingly, however, there are two studies of cypress-tupelo wetlands that provide similar results: Mitchell and Lancia (1990) found prothonotary warblers to be the most abundant bird species of their studies in South Carolina; the mean number of detections per 25 m radius 10-min point count ranged from 0.11 in a clearcut to 2.68 in the forest interior. Similar results were obtained by R. Sallabanks (unpubl. data) in a study of bottomland hardwood forests along the Roanoke River in North Carolina; the mean number of detections per unlimited radius 10-min point count ranged from 0.73 in levee forest to 2.21 in large patches of cypress-tupelo swamp forest. Again, the prothonotary warbler was the most common species censused. Whitcomb et al. (1981) found 40 males per sq km in their Maryland study area and Fowler and Fowler (1985) report 0.90 birds per 0.8 km segments of the Duck River in middle Tennessee.

TERRITORY SIZE: Territory size seems variable. Mean territory size was 1.48 ha in Michigan (Walkinshaw 1953), 0.50 ha in southern Illinois (Kleen 1973), and 0.48 ha in Tennessee (Petit 1989). Lefebvre et al. (1994) considered this species to be nonterritorial in winter in northeastern Venezuela mangroves. May roost communally during nonbreeding season (Warkentin and Morton 1995).

SITE FIDELITY: In Venezuela, exhibited some fidelity to wintering sites (Lefebvre et al. 1994).

Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: Y
Mobility and Migration Comments: Arrives in Tennessee early to mid-April. Arrives in Puerto Rico by October (sometimes as early as August), departs by end of March; appears to form small flocks before migrating north in spring (Raffaele 1983). Arrives in Costa Rica late August-October, departs by early to mid-March. Present in South America mainly September-March (Ridgely and Tudor 1989). (Stiles and Skutch 1989). Migrants uncommon on Gulf lowlands of Mexico.
Estuarine Habitat(s): Scrub-shrub wetland
Palustrine Habitat(s): FORESTED WETLAND, Riparian
Terrestrial Habitat(s): Forest - Conifer, Forest - Hardwood, Forest - Mixed, Shrubland/chaparral, Woodland - Conifer, Woodland - Hardwood, Woodland - Mixed
Special Habitat Factors: Standing snag/hollow tree
Habitat Comments: BREEDING: Mature deciduous floodplain, river, and swamp forests; wet lowland forest. Primary habitats are almost always near standing water; swamps that are somewhat open with scattered dead stumps are preferred. Bottomland forests and extensive willow thickets near lakes or ponds are also quite suitable. Requires dense underbrush along streambanks (Bushman and Therres 1988). Nests in cavity (natural, old woodpecker hole, bird box, etc.), in snag or living tree, often or always near or over water, at average height of 1.5-3 m (range 0.9-9.8 m); male selects territory, nest site, places some material before female's spring arrival. May be limited by the number of available nesting cavities. See Blem and Blem (1994) for information on composition and microclimate of nests in Virginia. This is the only eastern warbler that nests in tree cavities or other crannies (the only other cavity-nesting wood warbler (Parulidae) is Lucy's warbler [VERMIVORA LUCIAE]). Petit and Petit (1988) provided the first record of a prothonotary warbler using an open-cup nest built by another species (red-winged blackbird [AGELAIUS PHOENICEUS]).

Hamel et al. (1982) provided details on habitat use and suitability in Virginia, North Carolina, South Carolina, Georgia, and Florida. The following five vegetation types used by this species in five physiographic provinces (Piedmont, Sandhills, Inner Coastal Plain, Outer Coastal Plain, and Coastline) are listed in order of decreasing suitability: oak-gum-cypress is suitable at the sapling-poletimber successional stage, but optimal at the late successional sawtimber stage; both bay swamp-pocosin and elm-ash-cottonwood are only marginal at the sapling-poletimber stage and suitable at the sawtimber stage; southern mixed mesic hardwoods are only marginal in both the sapling-poletimber and sawtimber stages; and pond-pine pocosin is marginal at the late successional sawtimber stage only. In all cases, shrubs, midstory canopy and dead trees or limbs are used for all activities (foraging, nesting, perching, roosting, and singing), and snags must have a dbh of at least 15 cm.

Kahl et al. (1985) described habitat around song perches in Missouri as typically level terrain, with a small number of woody stems (< 2.5 cm dbh; < 2800 per ha, never > 4200), short ground vegetation (< 0.20 m, never > 0.36), and a high canopy (16-40 m, never < 12). Other important features included few dead stems (2.5-9.9 cm dbh; < 200 per ha, never > 250) and an intermediate subcanopy closure (30-80%, never < 10 or > 90).

NON-BREEDING: In migration, habitat includes dry woodland, scrub, thickets, and mangroves (AOU 1983). Commonly mist-netted in citrus groves in Belize (Mills 1989). Individuals were mist-netted in second-growth marshes, mangrove scrub, and seasonally flooded low forest in the northeastern coastal region of the Yucatan Peninsula (Ornat and Greenberg 1990). Usually occurs near water and most typically in mangroves in Puerto Rico (Raffaele 1989), though Faaborg and Arendt (1992) reported mist-net captures in dry forest in southwestern Puerto Rico. Records from Colombia include mangroves, freshwater swamps, coastal shrubs, and along the Rio Frio River in the edge of the foothills. The birds usually occur near water, but were also historically noted "in yellow, acacia-like trees on the border of stump land and dry forest, far from water" (Bent 1953).

Adult Food Habits: Invertivore
Immature Food Habits: Invertivore
Food Comments: Eats insects and spiders; forages on floating driftwood, half-submerged logs, tree trunks (Terres 1980); probes bark crevices, rolled leaves, and tangles, occasionally takes fruits or nectar (Stiles and Skutch 1989). See Petit and Petit (1988) and Petit et al. (1990) for detailed informarion on foraging ecology in Tennessee. In mangroves in Venezuela, ate various insects, mostly beetles, captured on foliage and twig substrates at low to moderate heights (Lefebvre et al. 1992). In Panama, diet includes fruits of the gumbo-limbo tree (BURSERA SIMURABA), a widespread tropical species common in advanced second-growth (Trainer and Will 1984).
Adult Phenology: Diurnal
Immature Phenology: Diurnal
Length: 14 centimeters
Weight: 17 grams
Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: Commonly breeds in the southeastern U.S. wherever there is suitable habitat: wooded wetlands, bottomland hardwood forests, and cypress swamps. Essential habitat requirements are water, shade, and older trees that provide nesting holes. The center of abundance is the South Atlantic Coastal Plain physiographic area. Little is known of winter habitat on the Caribbean slope of Central America, Colombia, and northern Venezuela. Most studies indicate a steady decline in populations of this neotropical migrant since the 1970s. The major threat is loss, degradation, and fragmentation of habitat as many wetlands are either permanently drained or flooded. Some mortality occurs through cowbird parasitism and competition with other species for nest sites. Stewardship needs include determination of the minimum area requirements for viable breeding populations and better censusing studies that penetrate this species' habitat.
Restoration Potential: Restoration currently is not needed. Where the species is of concern (e.g., Rhode Island), it is on the periphery of its range and has never been a common breeder. Should the species decline, restoration/recovery seems highly plausible if the correct habitat is provided.
Preserve Selection & Design Considerations: In the mid-Atlantic region, nesting generally occurs in forest tracts of at least 70-100 ha (Robbins 1979, 1980, Whitcomb et al. 1981).
Management Requirements: Among six species that clearly need management and/or monitoring attention overall in the southeastern U.S. (Hunter, unpubl. data). Older trees with cavities should be left intact. Timber harvesting practices (if any) should leave snags of decaying trees uncut, and should also leave small patches permanently uncut to provide older cavity trees in the future. To be suitable, cavity trees should have a dbh of at least 15 cm. Also, permanently uncut buffer zones should be created on both sides of streams to provide thick and shady vegetation along stream banks; these buffer zones are recommended to be at least 90 m wide (Bushman and Therres 1988).

Nest boxes readily are used (Fleming and Petit 1986; Petit et al. 1987; Petit 1989; Blem and Blem 1991, 1992, 1994). See Mitchell (1988) for specifications for the construction and placement of nest boxes. Reduced rates of predation and brood parasitism may occur in boxes compared with natural cavities. Preserves could therefore be more beneficial if filled with nest-boxes, especially in habitats where natural cavities are in short supply and/or house wrens are common. Placement of nest-boxes (over water or over land) can also affect competition from wrens. Prothonotary Warblers that nested over deep (20-100 cm) water in an inland swamp in northern Ohio were more likely to escape competition from wrens (Petit et al., unpubl. data). Predator-free habitat, with high, stable water levels and lower wren populations, appears to be more beneficial than are nest-boxes placed in low-quality habitat (Brush 1991). Because beavers may create suitable habitat, beaver reintroduction in some areas may beneficial, especially in regions like the Piedmont (H. LeGrand, Jr., pers. comm.).

Monitoring Requirements: Habitat is often difficult to access, so monitoring programs are difficult. BBS routes may be sufficient to track this species, although more intensive censusing studies (e.g., Mitchell and Lancia 1990, Sallabanks, unpubl. data) would be preferred. Annual surveys of known populations using point count censusing techniques are probably the best way to monitor this species (Mabey and Rice, pers. comms.). Studies that monitor breeding productivity provide critical information on population recruitment and dynamics; it is imperative that we establish the cause(s) of low population density and low breeding productivity.
Management Research Needs: Area requirements seem to be the least understood management parameter. Better estimates of minimum viable population size and impacts of cowbird parasitism and competition for nest-sites would also be useful. In addition, the effect of levees on water levels and warbler populations needs to be explored further (Brush 1991); in eastern Iowa, water level management may prove more beneficial than nest supplementation in maintaining or improving habitat.
Biological Research Needs: 1. Further refine distribution on the wintering grounds to identify focal areas for conservation. Conduct threat assessments on focal areas. More information based on natural cavity nests in habitats of different quality is needed to provide full understanding of the impacts of habitat degradation on breeding populations. Very little is known about the specific impacts of mangrove destruction or degradation on overwinter survival, and almost nothing is known about habitat use during migration. Such information is essential for any future conservation or management efforts (Petit, 1999).
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: 18Nov2014
NatureServe Conservation Status Factors Author: Dirrigl, F. J., Jr., G. Hammerson, and R. Sallabanks; Modified by Jue, Dean K.
Management Information Edition Date: 14Sep1993
Management Information Edition Author: SALLABANKS, R.; REVISIONS BY F.J. DIRRIGL, JR., G. HAMMERSON, AND D.W. MEHLMAN
Management Information Acknowledgments: An earlier version was improved by the careful reviews of Lisa Petit and Harry LeGrand, Jr. I also thank Lisa Petit and Charles Blem for providing reprints, preprints, and unpublished data. Unpublished manuscripts were also provided by Tim Brush and Rick Enser. Thanks also go to all Heritage biologists who responded to the ESA questionnaire: Alabama - Mark Bailey; Arkansas - Cindy Osborne; Connecticut - Dawn McKay; Delaware - Christopher Heckscher; Florida - Steve Jones; Georgia - Jon Ambrose; Iowa - Daryl Howell; Illinois - Vernon Kleen; Indiana - John Castrale and Michelle Martin; Kansas - Bill Busby; Kentucky - Brainard Palmer-Ball; Louisiana - Steve Shively; Massachusetts - Brad Blodget; Maryland - Lynn Davidson; Michigan - Mary Rabe; Minnesota - Mary Miller; Missouri - Brad Jacobs; Mississippi - Tom Mann; North Carolina - Harry LeGrand, Jr.; Nebraska - Mary Clausen; New Jersey - Rick Dutko; New York - Kathy Schneider; Ohio - Daniel Rice; Pennsylvania - Barb Barton; Rhode Island - Rick Enser; South Carolina - John Cely; Tennessee - Andrea Shea and Paul Hamel; Virginia - Sarah Mabey; and West Virginia - Barbara Sargent. Judith Soule of the Michigan Natural Features Inventory provided valuable advice on writing, Bruce Peterjohn of the Fish and Wildlife Service, Office of Migratory Bird Management, Patuxent Wildlife Research Center sent BBS data, and Melissa Morrison of The Nature Conservancy's Eastern Regional Office sent ELLINK reports.
Element Ecology & Life History Edition Date: 11Apr1996
Element Ecology & Life History Author(s): SALLABANKS, R., 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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