A Brief History of the Golden-winged Warbler: Part II

By Erin Cashion, Curator of Natural History

Author’s Note: Be sure to catch up with A Brief History of the Golden-winged Warbler Part I, here.

The Golden-winged Warbler was seemingly never common in Ohio. It had been variously described as a "transitory visitor" by Jared Kirtland in 1838; a "rare summer resident" by John Wheaton in 1879; and "irregular in distribution" by William Dawson in 1903.

A screenshot of two pages from a historic text. The left page has a colored drawing made from a specimen of a Golden-winged Warbler perched on a branch, and the facing page is text describing its natural history and a black-and-white photograph captioned “At the mouth of the Douda Run – “Morgan County’s Best.”
A screenshot of two pages from a historic text. The left page has a black-and-white photograph of a Golden-winged Warbler at its nest on the ground and text surrounding the photo. The facing page is text stating the species is “quite irregular in distribution” and also has the beginning of the species profile for Nashville Warbler.

Above screenshots are taken from Dawson, William L. 1903. The birds of Ohio; a complete scientific and popular description of the 320 species of birds found in the state. Columbus, Wheaton Pub. Co.

By the late 1800s, hybridizations between the Blue-winged Warbler (Vermivora cyanoptera) & Golden-winged Warbler (Vermivora chrysoptera) were well documented, but the academic debate on the origin of the Brewster’s and Lawrence’s Warbler would go on for another 30 years; meanwhile, the range of the Golden-wing was steadily shifting northward. And wherever the Golden-winged Warbler retreated, the Blue-winged Warbler did not take long to appear.

Two black-and-white photos, left photo shows a male Brewster's warbler in a temporary enclosure, right photo shows two very young, scraggly looking fledgling warbler offspring
A black-and-white photograph of a Golden-winged Warbler nest on the ground next to a raspberry thicket

Above photos taken from Carter, T. Donald, & Howland, R. H. 1923. A Brewster’s Warbler and His Brood. The Auk, 40(3), 423–430.

Narrative accounts in the 1870s describe the Golden-wings moving north into New York and the southern Atlantic states and becoming rare in southern Pennsylvania, where they had previously been abundant – although it’s unknown if this was a continuing northward range shift, or a new one.

In Connecticut, a survey of specimens and historical accounts1 indicates that neither species was common in the state before 1850, and there were no breeding records for Golden-wings until 1875. In the 1880s, there was a marked increase of Golden-wings, followed by an incursion of Blue-wings around a decade later, then increasing records of Brewster’s and Lawrence’s Warblers. By 1900, the hybrids were commonplace, and by 1980, Goldens were completely absent from southern Connecticut – having been supplanted entirely by Blue-wings and Lawrence’s Warblers.

Map showing current and historic Golden-winged Warbler range. From Streby, H. M., D. E. Andersen, and D. A. Buehler, 2016. Text reads: Figure 1.1. Golden-winged Warbler breeding distribution from ~1880 through 1980. Estimated distribution (hash marks) for 1880-1920 was derived largely from state ornithological accounts. Mid-1900s expansion in the north is shown in gray and contraction in the south and east in dark gray. Open circles represent additional early historical records that have previously been included in breeding-distribution maps, but which we regard as questionable (see text).

Map showing current and historic Golden-winged Warbler range. From Streby, H. M., D. E. Andersen, and D. A. Buehler, 2016

During the 20th century, the northerly population of Golden-wings went from nonexistent, to common, then to rare in northern Pennsylvania, while observations in Ontario and Manitoba increased. Meanwhile, the Appalachian golden-wing population suffered a 95% population decline from the 1960s onward, contracting into high elevation refugia on mountaintop balds2.

These exhaustive accounts of hybridization and range shifts do not, however, contain any conjecture why these species were interbreeding, and why the Golden-wing was being moving north (and uphill), seemingly being pushed by encroachment from the Blue-wing. The first mentions of Brewster’s (“White-winged”) and Lawrence’s Warbler in historical accounts are not made until 1874 – nearly a century after the parent species were first described. It begs the question: did the hybrids exist prior to that and go unnoticed by early naturalists, or were the two species not hybridizing in significant numbers until that time? After millions of years of being separated, what caused these two closely related species to reunite? What triggered the range shift for these two species?

A clue lies in what was happening to the landscape throughout much of the eastern United States during the 19th century.

By the early 1800s, the First Industrial Revolution was ramping up, and deforestation in the North American landscape was accelerating. Massive tracts of ancient forest habitat that separated these species were felled, first via selective thinning, then later via clearcutting, feeding iron furnaces and providing raw material for furniture, fences, homes, hearths, and railroads. Cleared forests were turned into farms, and areas unsuitable for farming were left raw and denuded.

In 1830, the Erie Canal opened a route between the Hudson River and Lake Erie, giving large, mechanized farms in western New York, Ohio and Indiana a competitive advantage over those further east. This encouraged farmers to abandon their smaller, rockier New England farms in favor of more arable properties to the west2. This deforestation, farm abandonment, and subsequent westward migration of settlers would continue over the next century. Harlan reported in The Birds of Ohio - 150 Years Later (1988)3 that precipitous population declines of several iconic species occurred between 1820 and 1870: the Passenger Pigeon (Ectopistes migratorius), Carolina Parakeet (Conuropsis carolinensis), and Greater Prairie Chicken (Tympanuchus cupido). By 1875, most of the original old growth forests were gone from Ohio, wetlands had been drained, and prairie habitat had been plowed and converted into farmland.

Around the turn of the century, successional habitat had begun to regenerate in the formerly farmed or forested landscapes – prime habitat for the Blue-winged & Golden-winged Warbler. It is not surprising that the first hybrids appear in the 1870s. The Golden-wings would arrive first, followed shortly by the Blue-wing. The latter, with its broader habitat preferences, was able to continue using the habitat after the Golden-wings no longer found it suitable; although they too would be pushed out as the trees aged. Without maintenance, early successional forests turn into woodland within 25 years or so, meaning this prime habitat was relatively short-lived.

This is the pattern throughout their ranges: the forests were cleared for the Industrial Revolution and the land was farmed for a century or so, then abandoned; as the forests began to grow back, first the Golden-wings moved in, followed by Blue-wings within 10-15 years. As the trees mature, the Goldens shift their range to the north and/or are absorbed into the Blue-wing population via hybridization. Once the Blue-wing arrives in Golden-wing territory, replacement of the Golden-wing by the Blue-wing occurs within 50 years1.

A screenshot of a historic text showing drawings of the Golden-winged Warbler, Blue-winged Warbler, and hybrid forms
The hybrid warblers shown in the color plate opposite are, from top to bottom:  F1 Brewster’s Warbler. Genotype WwSsPp. American Museum of Natural History Specimen No. 380136. New Haven, Connecticut, May 17, 1892. Back-cross Brewster’s Warbler. Genotype WWSSPp. American Museum of Natural History  specimen No. 392172. Bonilla, Costa Rica, September 27, 1920. Lawrence’s Warbler. Genotype wwsspp. American Museum of Natural History, L. C. Sanford collection No. 10846. Stratfield, Connecticut, May 19, 1917. Heterozygous Golden-winged Warbler. Genotype WwSsPp. Chicago Natural History Museum No. 148778. New Haven, Connecticut, May 11, 1900. Crossover-type Golden-winged Warbler. Genoytpe WWSspp. American Museum of Natural History No. 506277. Cumbre de Valencia, Venezuela, January 31, 1910. Crossover-type Blue-winged Warbler. Genotype wwSsPP. American Museum of Natural History No. 380055. New Haven, Connecticut, May 17, 1893.

The photo at left and caption above are taken from Parkes, Kenneth C. 1951. The genetics of the golden-winged X blue-winged warbler complex. Wilson Bulletin, 63(1), pp.5-15.

The first reports of hybridization in Ohio were made in the late 1920s by Lawrence E. Hicks. Hicks was one of the most prolific Ohio authors regarding songbird distribution and ecology in the first half of the 20th century, standing on the shoulders of Edwards, Bertram, and Wheaton. In his 1929 Some interesting Ohio records2 he reports “a pair of typical Brewster’s Warblers feeding young at Neotoma, Hocking County, on June 11 and 12, 1927. In 1928, only the male Brewster’s Warbler returned… Several Brewster’s Warblers, in addition to these records, were recorded near Columbus during the summer months of 1928.” This record of a nesting pair of Brewster’s Warblers would end up being one of only two reported cases of hybrids mating with each other. Also reported in this publication was a Lawrence’s warbler in Huron County, in what is now the Willard Marsh Wilderness area:

“The bird had the typical plumage of a Lawrence’s Warbler (the black throat and ear patches of the Golden-winged Warbler and the yellow crown and underparts and the two whitish wing bars of the Blue-winged Warbler) with one exception. Between the two white wing bars was a solid patch of gold, similar to the wing bar of the Golden-wing. The bird was singing a song indistinguishable from those of a half dozen Blue-winged Warblers singing nearby.”

In his 1933 report The Breeding Birds of Ashtabula County, Ohio5, he describes the Golden-winged Warbler as “decidedly local and rare everywhere in the state”, with single records reported in 17 counties, and a cluster of 8 within Ashtabula County. The Blue-wing was “rather local but ranging from rare to abundant. More frequent… in moist thickets and adjacent to boggy areas[The Brewster’s Warbler] appears to occur about as commonly as the Golden-winged Warbler... No evidence of nesting of this hybrid was obtained, in fact the behavior of individuals suggested in most cases that they were non-breeding birds. On three occasions a male golden-wing and a female blue-wing have been found feeding hybrid young.”

Which brings us to the specimen that started this whole thing: N 11948: the nest from a pairing between a female Blue-winged Warbler (Vermivora cyanoptera) and a male Golden-winged Warbler (Vermivora chrysoptera) in Ohio. Hicks writes:

On May 29, 1930, the writer found a nest with five eggs being incubated by a female blue-wing in southern Wayne Township. The next day, upon returning with Robert H. McCormick and Roscoe W. Franks, it was found that the eggs were hatched, several stills and movie pictures being taken of the hybrid young being fed by both parents at the same time, the female blue-wing and the male a typical Golden-winged Warbler.”

A screenshot of a historic text describing finding the hybrid nest (see text)
A still taken by Robert McCormick on May 30, 1930 of the female Blue-winged Warbler in the nest brooding the just-hatched young, and the male Golden-winged Warbler bringing food to the nest. You may be surprised to see that these warblers nest on the ground!

A still taken by Robert McCormick on May 30, 1930 of the female Blue-winged Warbler in the nest brooding the just-hatched young, and the male Golden-winged Warbler bringing food to the nest. You may be surprised to see that these warblers nest on the ground!

It is among the last recorded instances of the Golden-winged Warbler breeding in Ohio – and since the mate was a Blue-wing, the offspring raised in this nest were not Golden-wings, but Brewster’s Warblers.

In his 1935 Distribution of the breeding birds of Ohio7, Hicks reports the Golden-wing breeding in Vinton, Meigs, Scioto, Morgan, Monroe, Columbiana, Stark, Ashtabula, Trumbull, Huron, Williams, Fulton, Van Wert, Mercer, and Lucas Counties. The only area in which it was a common summer resident was in Oak Openings, in Lucas County. By contrast, the Blue-wing was noted as absent from large portions of the northwestern quarter of the state, where the Golden-wing was more likely to be found. Hybrids were reported in Ashtabula, Hocking, Franklin, Huron, Lorain, and Lucas counties.

A copy of Birds of the Toledo Area is at the Ohio History Center Archives & Library, its record is here.

By the mid-20th century it is concluded that human-caused habitat disturbance – i.e. the widespread deforestation and farm abandonment that took place in the 19th century – is likely at least one of the drivers behind hybridization. Ernst Mayr, one of the most well-known and prolific natural history authors of the 20th century, wrote of the species in 1942 in Systematics and the Origin of Species8: “it is probable that the hybridization is of recent date and caused by manmade habitat disturbances.”

But there was more to the story than the Blue-wing hybridizing them out of existence. Lou Campbell documented a dramatic decline of the Golden-winged Warbler between the 1930 and 1945 in Oak Openings, their last stronghold in the state. In his 1968 book Birds of the Toledo Area8, he reported 37 singing males in 1932; an average of 14 singing males from 1937-39; and 10 in 1945. From 1950 to the printing of the book in 1967, only 1 – 3 were seen each summer. He states, “Drainage, clearing of land for farming and home building, insect controls, and perhaps the loss of wilderness atmosphere have all combined to cause this decline.”

Taken from Streby et al. 2016. This map shows the distribution of phenotypic Golden-winged Warblers (GWWA), Blue-winged Warblers (BWWA), and zone of hybridization based on 1999–2005 data from the Golden-winged Warbler Atlas Project (GOWAP).

Taken from Streby et al. 2016. This map shows the distribution of phenotypic Golden-winged Warblers (GWWA), Blue-winged Warblers (BWWA), and zone of hybridization based on 1999–2005 data from the Golden-winged Warbler Atlas Project (GOWAP).

No breeding records for the Golden-winged Warbler have been reported in Ohio since the 1930s. All sightings since the 1980s have been of lone singing males in Oak Openings, Cuyahoga National Park, Glen Helen Preserve, and Lake Hope State Park. There were also no confirmed breeding records during the First (1982–1987) or the Second (2006–2011) Atlas of Breeding Birds in Ohio9. The most recent summer record occurred on June 2, 2023 in Columbiana County, Ohio – although there was no evidence of breeding, and many singing Blue-winged Warblers were present (see the record here: https://ebird.org/checklist/S140278759). The Blue-winged Warbler, having been reported in 80 of Ohio’s 88 counties by 19359, also experienced a significant range contraction during the latter half of the 20th century. Significantly fewer records were reported in the glaciated portion of the state during the Second Atlas as compared with the First Atlas10.

A copy of The second atlas of breeding birds in Ohio is at the Ohio History Center Archives & Library, its record is here.

It appears separation between these two species is now no longer possible. Brought into contact by human-caused disturbance, the future of the imperiled Golden-winged Warbler is now uncertain. Early successional habitat is now extremely scarce in Ohio, and indeed in much of the Eastern United States. Marshes, wetlands, bogs, and the otherwise shrubby, wet habitats favored by the Golden-wing (and many other species!) have long since been drained, tiled, and turned into agriculture. Early successional habitats also suffer from a PR problem: the tall flowering forbs, swaying grasses, and dense shrubby growth that characterize them are viewed as unaesthetic, something to be mowed into homogeneity. But early successional forests are not only important breeding habitat for the Golden-winged Warbler; these structurally complex habitats with ample food sources and cover are also heavily used by other songbird species during fall migration11.

A female Golden-winged Warbler, Vermivora chrysoptera. Photo by Chrissy McClaren and Any Reago (flickr user Wildreturn). https://creativecommons.org/licenses/by/4.0/

A female Golden-winged Warbler, Vermivora chrysoptera. Photo by Chrissy McClaren and Any Reago (flickr user Wildreturn). https://creativecommons.org/licenses/by/4.0/

The future of the Golden-winged Warbler is uncertain. If management of early successional habitats is not prioritized, the Golden-winged Warbler may be driven to extinction in the relatively near future throughout its range – both by a lack of breeding habitat, and by genetic swamping by its close relative, the Blue-winged Warbler. However, if sufficient efforts are made to restore and maintain these once-scarce habitats, it may be possible to hear the raspy buzz of the Golden-winged Warbler in Ohio once again.

References

[1] Gill, Frank B. 1980. "Historical aspects of hybridization between Blue-winged and Golden-winged warblers." The Auk 97.1: 1-18. https://sora.unm.edu/sites/default/files/journals/auk/v097n01/p0001-p0018.pdf

[2] Kingsley, N. P. 1974. The timber resources of southern New England. USDA Forest Service Resource Bulletin NE-36. https://archive.org/details/timberresourceso36king

[3] Streby, H. M., D. E. Andersen, and D. A. Buehler. 2016. Golden-winged warbler ecology, conservation, and habitat management. Studies in Avian Biology, Cooper Ornithological Society, CRC Press/ Taylor & Francis. ISB no. 13:978-1-4822-4068-9. https://conservancy.umn.edu/items/d4dca0dc-7c66-4682-ab06-25ba0192c2d4

[4] Harlan, Robert. 1988. The Birds of Ohio - 150 Years Later. Ohio Cardinal: Vol. 11:3, Article 1. https://digitalcommons.usf.edu/ohio_cardinal/vol11/iss3/1

[5] Hicks, Lawrence E. 1929. Some Interesting Ohio Records. Wilson Bulletin: Vol. 41:1, 43-44. https://www.jstor.org/stable/4155716?seq=1

[6] Hicks, Lawrence E. 1933. The Breeding Birds of Ashtabula County, Ohio. The Wilson Bulletin, 45(4), 168–195. http://www.jstor.org/stable/4156234

[7] Hicks, Lawrence E. 1935. Distribution of the breeding birds of Ohio. Columbus, Ohio: Ohio State University Press. https://babel.hathitrust.org/cgi/pt?id=mdp.39015068570509

[8] Campbell, Louis W. 1968. Birds of the Toledo Area. The Blade.

[9] Mayr, Ernst. 1942. Systematics And The Origin Of Species. Columbia University Press, New York, p. 262. https://archive.org/details/in.ernet.dli.2015.20284/page/n277/mode/2up

[10] Rodewald, P. G., Shumar, M. B., Boone, A. T., Slager, D. L., & McCormac, J. 2016. The second atlas of breeding birds in Ohio. The Pennsylvania State University Press.

[11] Cashion, Erin. 2011. Avian use of riparian habitats and the conservation reserve program: migratory stopover in agroecosystems. MS thesis. The Ohio State University. https://etd.ohiolink.edu/acprod/odb_etd/ws/send_file/send?accession=osu1307904873&disposition=inline

 

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