A large (6 inches) wood warbler, the Northern Waterthrush is most easily identified by its brown back and wings, yellowish breast streaked with brown, and yellowish eye stripe. This species is physically similar to the related Louisiana Waterthrush (Parkesia motacilla), although that species is typically paler and less streaked below and on the face. Male and female Northern Waterthrushes are similar to one another in all seasons. The Northern Waterthrush breeds across Alaska, Canada, and the northern tier of the United States. In winter, this species is primarily found in the southern half of Mexico, Central America, and the West Indies. Small numbers of Northern Waterthrushes winter in south Florida, mainly south of Miami. Northern Waterthrushes breed in a variety of cool woodland habitats along the edges of shallow bodies of water, including streams, ponds, and bogs. In winter, this species is found in wetland portions of humid tropical forests as well as in coastal mangrove forests. Unusually for a warbler, Northern Waterthrushes primarily eat aquatic invertebrates, including insects and larvae, although this species will also eat terrestrial insects, snails, and small crustaceans during the winter. Along bodies of water in appropriate habitat, Northern Waterthrushes may be seen walking on the shoreline or wading in shallow water while foraging for food. This species’ characteristic tail wagging behavior, in which the rear half of the body is flicked up and down almost constantly while the bird is in motion, is highly unusual among wood warblers. Northern Waterthrushes are primarily active during the day, but, like many migratory songbirds, this species migrates at night.
- Eaton, Stephen W. 1995. Northern Waterthrush (Parkesia noveboracensis), The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/species/182
- Peterson, Roger Tory. Birds of Eastern and Central North America. Boston: Houghton Mifflin, 1980. Print.
- eBird Range Map - Northern Waterthrush. eBird. Cornell Lab of Ornithology, N.d. Web. 20 July 2012. http://ebird.org/ebird/map/norwat.
- Seiurus noveboracensis. Xeno-canto. Xeno-canto Foundation, n.d. Web. 20 July 2012. http://xeno-canto.org/browse.php?query=Seiurus+noveboracensis.
- Northern Waterthrush (Seiurus noveboracensis). The Internet Bird Collection. Lynx Edicions, n.d. Web. 20 July 2012. http://ibc.lynxeds.com/species/northern-waterthrush-seiurus-noveboracensis.
occurs (regularly, as a native taxon) in multiple nations
Regularity: Regularly occurring
Type of Residency: Breeding
Regularity: Regularly occurring
Type of Residency: Breeding
Global Range: BREEDING: western and north-central Alaska and northwestern Mackenzie to Labrador and Newfoundland, south to southeastern British Columbia, Washington, Idaho, North Dakota, Great Lakes, eastern West Virginia, northwestern Virginia, and Massachusetts (Eaton 1995, AOU 1998). NON-BREEDING: southern Baja California, southern Sinaloa, San Luis Potosi, northern Veracruz, and southern Florida south through Mexico (including Yucatan peninsula), throughout Central America to Colombia, Ecuador, Venezuela, northern Brazil, northeastern Peru, and Surinam; also abundant throughout the West Indies (Raffaele 1983, Pashley 1988a, Pashley 1988b, Pashley and Hamilton 1990, Eaton 1995).
- Clements, J. F., T. S. Schulenberg, M. J. Iliff, B.L. Sullivan, C. L. Wood, and D. Roberson. 2012. The eBird/Clements checklist of birds of the world: Version 6.7. Downloaded from http://www.birds.cornell.edu/clementschecklist/downloadable-clements-checklist
In the breeding season, the Northern Waterthrush is found in a belt stretching from north central Alaska, east across all of the Canadian provinces. In the winter season, the species is found in the tropical mangroves of Central and South America.
Biogeographic Regions: nearctic (Native ); neotropical (Native )
The Northern Waterthrush is a large, ground walking warbler with a brown back and a white or yellowish streaked breast. The breast, sides, and flanks are streaked with a dark olive or black. There is an olive-colored triangular spot in the front of the eye and a crescent shaped mark on the lower eyelid. The throat is also covered with small triangular marks.
Average mass: 18.7 g.
Average basal metabolic rate: 0.28391 W.
Length: 15 cm
Weight: 18 grams
Catalog Number: USNM 381382
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Birds
Sex/Stage: Male; Adult
Preparation: Skin: Whole
Collector(s): H. Peters & T. Burleigh
Year Collected: 1945
Locality: Topsail, Saint John'S, Newfoundland, Canada, North America
- Type: Burleigh & Peters, H. S. June 16, 1948. Proc. Biol. Soc. Washington. 61: 120.
Catalog Number: USNM 370456
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Birds
Sex/Stage: unknown; Adult
Preparation: Skin: Whole
Collector(s): S. Williston
Year Collected: 1878
Locality: Como, Carbon, Wyoming, United States, North America
- Type: Ridgway. April? 1880. Proc. U.S. Nat. Mus. 3: 12.
Comments: BREEDING: Breeds in cool, wooded swamps, ponds and slow-moving rivers; thickets of bogs, and rivers bordered with willow (SALIX) and alder (ALNUS, Godfrey 1986, Peck and James 1987). Regional habitats differ slightly. Throughout Canada and Alaska, nests primarily in spruce (PICEA) bogs, along alder-and willow-bordered rivers; also along lakes, swamps, and wet woodlands (Godfrey 1986). However, on islands off Newfoundland, known to nest in areas without standing water and where understory is less dense than on mainland (Vassollo et al.1982).
In New York state, breeds in hardwood swamps dominated by Red Maple (ACER RUBRUM) on the Great Lakes Plain, in Eastern Hemlock (TSUGA CANADENSIS)-northern hardwood swamps on the Allegheny Plateau, and in spruce-tamarack (LARIX)-balsam (ABIES) swamp valleys and uplands of the Adirondacks and Tug Hill Plateau (Eaton 1988). In Pennsylvania, found in RHODODENDRON swamps and a variety of wooded wetland types (Gross 1992). In Massachusetts, nests in locally in red maple swamps and white cedar and red maple swamps (Viet and Petersen 1993). In West Virginia, nests along rhododendron-bordered mountain streams, in spruce swamps and northern mixed forest to beech (FAGUS)-maple (ACER) forest (Brooks 1944).
Where sympatric with Louisiana Waterthrush, nests in areas with more forbs and ferns, with significantly more moss cover, hummocks, and conifers and with a higher density of shrubs; significantly more Eastern Hemlock and alder in Northern Waterthrush territories (Craig 1985).
NONBREEDING: Found mainly in damp tropical lowland forest, edges of pools and streams, mainly below 1,500 m. Mangroves (RHIZOPHORA, AVICENNIA, LABUNCULARIA) provide key habitat throughout much of range (Stotz et al. 1996, Bond 1971, Wetmore et al. 1984, Binford 1989, Lefebvre et al. 1992, Wunderle and Waide 1993). In Costa Rica, also found in open second growth or at wet spots in trails or roads (Stiles and Skutch 1989 Blake and Loiselle 1992). In northeast Nicaragua, in rain forest adjacent to pine habitat (Howell 1971). Tends to avoid disturbance, but may do well in second-growth tropical forest, edges, or woodlots (Ehrlich, et al. 1988).
Throughout its winter range found mainly below 1500 m (Curson et al. 1994). Highest recorded elevation in Columbia 3000m (Hilty and Brown 1986). In Costa Rica, ranges from lowlands to 1500 m, rarely higher, on both slopes, often extremely abundant in September along Caribbean coast; most numerous in Caribbean lowland and mangroves along the Pacific coast (Stiles and Skutch 1989). In Belize, most numerous in mangroves and gallery forest (Eaton 1995); occurs sparingly in cropland of cacao, rice, and citrus fruits (Robbins et al. 1992). In the West Indies, most often near the border of standing water, primarily saline and brackish, in or near mangroves and coastal scrub forest (Raffaele et al. 1998).
MIGRATION: Prefers damp woodlands with standing water, thick cover along streams, in marshes, and by stagnant pools, but is also found on lawns and in hedgerows and thickets (Winkler et al. 1992).
Habitat and Ecology
The Northern Waterthrush prefers cool, dark, wooded swamps, thickets of bogs, margins of northern lakes, and willow and alder bordered rivers. During the spring and fall migration, the bird can be found in thick cover along streams, in marshes, and by stagnant pools.
Terrestrial Biomes: forest ; rainforest
Aquatic Biomes: lakes and ponds; rivers and streams
Non-Migrant: No. All populations of this species make significant seasonal migrations.
Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).
Locally Migrant: Yes. At least some populations of this species make annual migrations of over 200 km.
A neotropical migrant, traveling long distances nocturnally (Winkler et al. 1992). Migrates annually between breeding grounds in North America and wintering areas in West Indies and Central and South America, flying across or around Gulf of Mexico; a trans-gulf, circum-gulf, and trans-Caribbean migrant (Eaton 1995). Cannot carry enough fat to complete spring or fall migrations in one direct flight (est. 2,500 km) and must stop and eat (Winkler et al. 1992); migratory stopover habitat is especially important to this species.
Present in South America mainly September-April (Ridgely and Tudor 1989). Arrives in Puerto Rico and the Virgin Islands in September, present through April (Raffaele 1983). Arrives in Costa Rica mid- to late-August, departs by mid-May; often abundant in September (Stiles and Skutch 1989).
Comments: On breeding grounds, eats various small invertebrates, primarily from muddy ground but also wades in shallow pools, gleans from foliage or soggy fallen leaves, and occasionally catches flying insects (Terres 1980, Lack 1976, Rappole and Warner 1980, Craig 1984). Takes larval and adult insects, spiders and snails. In the north, where associated with moving water, probably feeds on stoneflies (Plecoptera). After leaf emergence in spring, feeds extensively on Lepidoptera larvae (Eaton 1957, Craig 1987, Eaton 1995) In S. Carolina, known to have taken small minnows (Wayne 1910).
On non-breeding grounds, forages in mangroves, perching on fallen trees or pneumatophores, picking prey from substrates or near water surface, up to 3 m from ground but usually below 1 m (Lefebvre et al. 1992). Diet includes beetles, ants (Hymenoptera), flies, insect larvae, snails and decapod crustaceans found at water surface, on the ground, on fallen trees, or occasionally in low foliage. Typically forages alone, but sometimes in small groups probably only during migration when birds are immediately concerned with feeding (Schwartz 1964). In Cuba, small snails, small clams (Pelecypodia), snout beetles (Rhynchophora), small spiders and ants (Eaton 1995). Feeds on a greater variety of prey during migration.
During the breeding season, the basic diet consists of larval and adult insects, spiders, and snails. After leaf emergence in the spring, the bird feeds on primarily on butterfly larvae. On the winter grounds, the bird consumes a greater variety of food, adding minnows and decapod crustaceans to its diet. The Northern Waterthrush forages alone using such tactics as twig gleaning, flycatching, hovering, chasing, and a lot of pecking. Microhabitats for foraging include water, ground, foilage, and air. Before leaf emergence, the Waterthrush typically spends about 75% of the time feeding in water, alternating between wading and walking along logs, on branches, and the water's edge.
Territorial throughout the year. Thought to defend non-breeding foraging areas against intraspecific intrusion, occasionally violently. Mean territory size on breeding grounds from 0.5-1.0 ha by location and is similar on wintering grounds (Eaton 1995, Curson et al. 1994). Some indication that individuals may show changing preferences for habitat throughout non-breeding season despite other studies showing strong winter territoriality. Lefebvre et al. (1994) considered this species to be non-territorial in winter in northeastern Venezuela mangroves. Arrivals in Venezuela near end of rainy season occupy higher slopes, descending to humid lowlands in the dry season (Schwartz 1964). In northern Colombia, inhabits thornscrub in October but disappears in November as leaves wilt; unrecorded there in spring (Russell 1980). In addition, birds occupying coastal mangroves in Panama may migrate between habitat types mid-winter in response to prey availability. Abundance of Panamanian birds followed patterns of arthropod abundance and increased rainfall; species was more abundant in Pacific mangrove forests during the first part of the wintering period and more abundant in the Caribbean mangroves during the second part of the wintering season when rainfall and arthropod prey items increase there (Lefebvre and Poulin 1996).
Life History and Behavior
Status: wild: 107 months.
Lifespan, longevity, and ageing
Essentially monogamous. Pair bond maintained from shortly after male arrives to 3-4 days after successful fledging. Favors nest sites in cavities of root systems of wind-blown trees in wooded swamps, or on sides of fern clumps or under cover on the banks of lakes or rivers. Nest typically hidden from above (Eaton 1995, Baicich and Harrison 1997). Nest a bowl of moss and liverwort gametophytes with a few leaves on the outside, lined with grass stems, twigs or pine needles, moss sporophytes or small rootlets and hair. May have an entranceway of leaves (Eaton 1995, Baicich and Harrison 1997).
Clutch size four to five eggs, sometimes three to six. Distinctly smaller than cowbird eggs. Female incubates and will lure potential predators away from nest. (Eaton 1995). Eggs are laid in late May-June. Young are altricial, brooded by female until day five. Both parents feed young. Departure from nest at day nine. Young unable to fly and hide for 2-3 days under dense vegetation (Baicich and Harrison 1997). Parents split brood for feeding. One brood per season (Eaton 1995).
Pair bonding begins immediately after the female arrives on the breeding site. Males perch in trees that are standing near water while females are below usually feeding along the edge of the water. Males vibrate their wings and raise their crown feathers and sing. The female may answer with a chink. The pair bond is broken shortly after successful fledging. The male selects the general area of the nesting site, but the female chooses the actual nest site. Nests are usually in the cavity of a root system of wind-blown trees in a wooded swamp, on the side of a fern clump or under cover along the banks of a lake or a river. There is typically covering above the nest and an opening to one side. The exterior of the nest is mainly moss and liverwort gametophytes. The interior of the nest cup is constructed with grass stems, twigs, or pine needles, and then lined with the hair of deer, caribou, cow, and rabbit. Average clutch is composed of four white ovate eggs spotted with browns and greys. The female is solely responsible for incubation, which lasts 12 days. Both parents, however, share feeding responsibilities. After four to five weeks the chicks begin feeding themselves.
Average eggs per season: 4.
Molecular Biology and Genetics
Barcode data: Seiurus noveboracensis
There are 26 barcode sequences available from BOLD and GenBank. Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species. See the BOLD taxonomy browser for more complete information about this specimen and other sequences.
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Download FASTA File
Statistics of barcoding coverage: Seiurus noveboracensis
Public Records: 27
Specimens with Barcodes: 36
Species With Barcodes: 1
National NatureServe Conservation Status
Rounded National Status Rank: N5B - Secure
Rounded National Status Rank: N5B - Secure
NatureServe Conservation Status
Rounded Global Status Rank: G5 - Secure
IUCN Red List Assessment
Red List Category
Red List Criteria
Degradation of habitat and pesticides are the biggest threats facing the species. Northern Waterthrush populations, however, have managed to remain stable despite these threats. Drainage of swamps for agriculture and wetland development into ponds or lakes may reduce breeding habitat. Pesticides are also affecting the Northern Waterthrush. Aerial spraying for the spruce budworm can kill the birds directly or reduce the biomass of their prey.
US Migratory Bird Act: protected
US Federal List: no special status
CITES: no special status
State of Michigan List: no special status
IUCN Red List of Threatened Species: least concern
Comments: HABITAT LOSS: Habitat appears relatively secure throughout Canada, the majority of breeding range, but is increasingly threatened on wintering grounds. As human populations in the West Indies, Mexico, Central America, and northern South America are increasing, felling of mangrove forests will increase in response to demands for fuel, food and space (Eaton 1995). Current estimates show that only 21% of land area remains forested (Wunderle and Waide 1994). The mangroves of Ecuador and Peru have been decimated for fuel and paper making and more recently for shrimp culture impoundments. Mangroves of Central America and the Caribbean also are threatened by such activities (Terborgh 1989). Throughout range, drainage of swamps for agriculture and wetlands development into ponds or lakes also destroy and degrade habitat on breeding and non-breeding grounds (Eaton 1988, Eaton 1995, Hull 1991, Gross 1992). Forest fragmentation and activities that cause reductions in forest canopy cover or negatively impact aquatic insect communities are also a threat (Brown 1999). PESTICIDES AND CONTAMINANTS: Wetlands frequently concentrate environmental contaminants and pesticides, putting species that use these habitats at high potential risk for mortality from ingestion (Rappole et al. 1983). In addition, northern forests on breeding grounds are often sprayed to control spruce budworm outbreaks. This could affect Northern Waterthrush populations by reducing insect prey base and directly killing birds (Eaton 1995). Potential impacts to prey base due to acid precipitation caused by powerplants in some regions. COLLISIONS: One record of a large kill due to collision with TV towers where 517 birds were killed (Robbins 1991). HURRICANES: Hurricanes may reduce local wintering populations of neotropical migrants in the West Indies (Holmes 1994; J. Confer and R. Holmes, unpubl. data). PREDATION: Young and eggs may experience high predation rates (Eaton 1957). PARASITISM: Relatively free of parasitism by the Brown-headed Cowbird (MOLOTHRUS ATER) compared with the Louisiana Waterthrush and other North American warblers (Eaton 1995).
Restoration Potential: Not currently in need of restoration. Populations throughout much of North America appear stable. However, as noted, the most serious threats may come on wintering areas.
Preserve Selection and Design Considerations: Essential habitat includes thickets along edges of wooded swamps, montane woodland with bogs and pools of standing water. Fallen trees with exposed root masses are preferred nest sites. Absent in highly fragmented landscapes. Wetlands altered by dredging and sedimentation and pollution are suboptimal.
Management Requirements: Ability to maintain populations is closely tied with habitat preservation; threats appear more severe to non-breeding habitat. On breeding grounds, wooded streambanks should be protected. Throughout non-breeding grounds, mangroves and riparian/gallery forest are closely associated with this species, and successful management will focus on coastal and riparian forest preservation.
This and many other migratory species that overwinter in South America are associated with relatively tall, moist/wet forests, as well as some moderately disturbed sites, gaps and forest edges. Maintaining such forests is an extremely important part of any management strategy for this and a host of other Neotropical migrants; conversion to agricultural lands or pasture would remove the critical structural components of these habitats (Petit et al. 1996). Mangroves in Caribbean and coastal areas of Venezuela provide crucial wintering areas for this species and need protection (Lefebvre et al. 1992, Wunderle and Waide 1993).
Second-growth tropical forest edge and woodlots may provide sufficient habitat in some locations. However, deforestation in the tropics only rarely results in such habitats, but more often in vast expanses of overgrazed pastures, canefields and other inhospitable habitats (Ehrlich et al.1988).
Also, there are indications that fragmentation of tropical forests may lead to increased mortality for migrants occupying those patches. Until further research proves otherwise, limiting fragmentation and increasing forest connectivity should be guidelines for forest management in the tropics as well as in the temperate forests of North America (Petit et al. 1996).
Throughout the densely populated Caribbean islands, successful conservation depends in large part on identifying and implementing land-use practices that are compatible with both economic goals and preservation of natural vegetation types (Petit et al. 1996). Arendt (1992), in collaboration with other bird experts from the region, outlined four major requirements to achieve that goal in the Caribbean: 1) establishment of protected reserves; 2) restoration of degraded habitats; 3) legislation and education, 4) and ecological research (Petit et al. 1996, also see Wunderle and Waide 1994).
Management Research Needs: Further research is needed on breeding biology. Details of demography and population dynamics including factors affecting survivorship are poorly understood. In order to better assess conservation needs and design effective approaches more data are needed on the status, habits, and ecology of this species on its wintering grounds (Eaton 1995).
The effect of habitat fragmentation and loss of connectivity in tropical environments is poorly understood. Further investigation of this question is of immediate concern because of the rapidly increasing human populations throughout Central and South America and the Caribbean.
Biological Research Needs: None.
Relevance to Humans and Ecosystems
Economic Importance for Humans: Negative
Economic Importance for Humans: Positive
Feed on insects that are regarded as pests to humans.
Stewardship Overview: Locally common breeder in montane riparian woodlands and wooded bogs throughout Canada, Alaska, and the northeastern U.S. Threatened by destruction and degradation of mangrove swamps throughout non-breeding range, wetland destruction and degradation in North America. Collisions with communication towers also an increasing threat. Successful management of this species will require protection of remaining mangrove forests (nonbreeding) and spruce bogs and other wooded riparian habitat (breeding); restoration of degraded habitats is also a priority. Alternatives to the use of pesticides and further construction of tall communication towers should be investigated.
The Northern Waterthrush (Parkesia noveboracensis) is one of the larger New World warblers. It breeds in the northern part of North America in Canada and the northern United States including Alaska. This bird is migratory, wintering in Central America, the West Indies and Florida, as well as in Venezuela, Colombia, and Ecuador. It is a very rare vagrant to other South American countries and to western Europe.
The Northern Waterthrush is a large New World warbler (and not a thrush, despite the name). It has a length of 12–15 cm (4.7–5.9 in), wingspan of 21–24 cm (8.3–9.4 in) and weighs between 13 and 25 g (0.46 and 0.88 oz) Among standard measurements, the wing chord is 6.8 to 8.2 cm (2.7 to 3.2 in), the tail is 4.5 to 5.7 cm (1.8 to 2.2 in), the bill is 1.1 to 1.2 cm (0.43 to 0.47 in) and the tarsus is 1.9 to 2.3 cm (0.75 to 0.91 in). On the head, the crown is brown with a white supercilium. The bill is pointed and dark. The throat is lightly streaked brown to black with heavier streaking continuing onto the breast and flanks. The back is evenly brown. Sexes are morphologically similar. Young birds have buff, rather than white underparts.
The only species bird watchers confuse with the Northern Waterthrush is the closely related Louisiana Waterthrush (Parkesia motacilla), which has buff flanks, a buff undertail, and bright pink legs. The Louisiana Waterthrush also has a whiter throat with fewer streaks.
Both waterthrush species walk rather than hop, and seem to teeter, since they bob their rear ends as they move along.
On the wintering grounds in Puerto Rico, Northern Waterthrushes leave daytime foraging areas and fly up to 2 km (1.2 mi) to nighttime roosts. The roosts are often located in red mangrove habitats.
Northern Waterthrushes winter in 4 main habitats in Puerto Rico: white mangrove, red mangrove, black mangrove, and scrub. Males, which are larger and migrate earlier in spring, prefer to winter in white mangrove, and are able to maintain or gain weight through the winter. Females winter in the other drier and less food-rich habitats. Waterthrushes wintering red and black mangrove can maintain body weight through the winter but lose weight in scrub.
The breeding habitat of the Northern Waterthrush is wet woodlands near water. It nests in a stump or among tree roots where it lays three to six eggs, cream- or buff-colored, with brown and gray spots. These eggs are laid in a cup nest constructed of leaves, bark strips, and rootlets.
- BirdLife International (2012). "Parkesia noveboracensis". IUCN Red List of Threatened Species. Version 2013.2. International Union for Conservation of Nature. Retrieved 26 November 2013.
- R. Terry Chesser, Richard C. Banks, F. Keith Barker, Carla Cicero, Jon L. Dunn, Andrew W. Kratter, Kirby J. Lovette, Pamela C. Ramussen, J. V. Remsen, Jr., James D. Rising, Douglas F. Stotz, and Kevin Winker (2010). "Fifty-First Supplement to the American Ornithologists' Union Check-List of North American Birds". The Auk 127 (3): 726–744. doi:10.1525/auk.2010.127.4.966.
- Eaton, S. W. (1995). "Northern Waterthrush (Parkesia noveboracensis), The Birds of North America Online". Ithaca, NY: Cornell Lab of Ornithology. Retrieved 2008-04-16.
- "FieldGuides: Northern Waterthrush Species Detail". eNature.com. Retrieved 2012-08-24.
- Curson, Jon; Quinn, David; Beadle, David (1994). New World Warblers: An Identification Guide. Christopher Helm Publishers. ISBN 0-7136-3932-6.
- Smith, J. A. M., Reitsma, L. R., Rockwood, L. L. & Marra, P. P. (2008). "Roosting behavior of a Neotropical migrant songbird, the northern waterthrush Seiurus noveboracensis, during the non-breeding season". Journal of Avian Biology 39: 460–465. doi:10.1111/j.0908-8857.2008.04227.x.
Names and Taxonomy
Comments: Formerly (AOU 1983, 1998) placed in the genus Seiurus; transferred to Parkesia by AOU (2010).