The Re-Establishment of Pileated Woodpeckers in New York City Following Nearly Two Centuries of Extirpation

Abstract

Dryocopus pileatus (Pileated Woodpecker) is an endemic North American woodpecker that, following widespread declines during the mid-to-late 19th century, re-bounded and has become relatively common resident throughout its range. In New York City, Pileated Woodpeckers were breeding residents until the early to mid-1800s, but were extirpated due to the city’s rapid urbanization. Since the 1930s, however, the number of re-ports of Pileated Woodpeckers has increased dramatically in the area. Here we document the re-establishment of Pileated Woodpeckers in New York City within 2 old-growth greens-paces on Staten Island from 2015 to 2019. We further describe and validate a previously unpublished breeding attempt on Staten Island from 1989. The establishment of Pileated Woodpeckers in New York City is reflective of their ongoing colonization of urban and suburban areas throughout North America, and highlights the importance of unfragmented greenspaces within urban and suburban areas.

Full text

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J.R. Ramírez-Garofalo, S.R. Curley, A.V. Ciancimino, R.V. Matarazzo, E.W. Johnson, and R.R. Veit
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NORTHEASTERN NATURALIST
2020 27(4):803–816
The Re-establishment of Pileated Woodpeckers in New York
City Following Nearly Two Centuries of Extirpation
José R. Ramírez-Garofalo1,*, Shannon R. Curley1, Anthony V. Ciancimino1,
Raymond V. Matarazzo2, Edward W. Johnson2, and Richard R. Veit1,3
Abstract - Dryocopus pileatus (Pileated Woodpecker) is an endemic North American
woodpecker that, following widespread declines during the mid-to-late 19th century, re-
bounded and has become relatively common resident throughout its range. In New York
City, Pileated Woodpeckers were breeding residents until the early to mid-1800s, but were
extirpated due to the city’s rapid urbanization. Since the 1930s, however, the number of re-
ports of Pileated Woodpeckers has increased dramatically in the area. Here we document the
re-establishment of Pileated Woodpeckers in New York City within 2 old-growth greens-
paces on Staten Island from 2015 to 2019. We further describe and validate a previously
unpublished breeding attempt on Staten Island from 1989. The establishment of Pileated
Woodpeckers in New York City is reective of their ongoing colonization of urban and
suburban areas throughout North America, and highlights the importance of unfragmented
greenspaces within urban and suburban areas.
Introduction
Habitat loss and fragmentation, often as a result of urbanization (Aronson et
al. 2014) or development for agriculture, has caused the decline and extinction of
species worldwide (Brooks et al. 2002, Fahrig 2003). As a response, conservation
initiatives, government mandates, and local charters have been created to prevent
further loss of biodiversity (Ancrenaz et al. 2007, Tabarelli et al. 2005). At a broad
scale, many species have benetted from these practices, and species have been
conserved through land-preservation measures such as the establishment of na-
tional parks and greenspaces in cities and towns (Aronson et al. 2017).
In northeastern North America, forests were historically subject to mass harvest
and clearing for both agricultural use and the construction of human settlement
(Sanderson 2013, Thompson et al. 2013). During the last century, however, much of
the region has been subject to a period of forest regrowth (Thompson et al. 2013).
As a result of reforestation, some forest-interior specialists like Dryocopus pilea-
tus L. (Pileated Woodpecker) have become conservation success stories. Pileated
Woodpeckers are endemic to the North American continent. They typically have
large home-range sizes (>400 ha; Bull and Houlthausen 1993, Bull and Meslow
1977, Mellen et al. 1992, Renken and Wiggers 1989) and extensively use snags
(standing dead trees) for foraging, roosting, and nesting (Hoyt 1957). Their large
1Biology Department, The College of Staten Island, City University of New York, Staten
Island, NY 10314. 2The Staten Island Museum, Staten Island, NY 10301. 3Biology De-
partment, The Graduate Center, CUNY, New York, NY 10010. *Corresponding author -
jose.ramirez.garofalo@gmail.com.
Manuscript Editor: Susan Smith Pagano

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nest cavities are used by secondary-cavity–nesting species such as Aix sponsa L.
(Wood Duck) and Martes americana (Turton) (American Marten), making them
a keystone species throughout their range (Aubry and Raley 2002, Bonar 2000).
During the latter half of the 19th century, Pileated Woodpeckers underwent drastic
declines across North America. These declines have been mainly attributed to the
widespread clearing of forest for agricultural purposes in the 19th century (Bull
and Jackson 2011), with hunting playing a secondary role, particularly where sport
shooting was popular (e.g., Wisconsin; Stoddard 1947).
Pileated Woodpecker populations began to recover at the turn of the 20th century
when state, federal, and provincial bird protections were established, and forests
began to regrow (Bull and Jackson 2011). However, the species remained largely
absent from the coastal regions, and were mainly conned to heavily forested ar-
eas of the interior, such as the Adirondack Mountains in central New York and the
Berkshires in western Massachusetts (Eaton 1914, Forbush 1927). By the 1920s,
Pileated Woodpeckers began to habituate to suburban areas and as a result, became
increasingly common at backyard bird feeders throughout the northeast (Hoyt
1957) and in secondary-growth forests (Bull and Jackson 2011, Hoyt 1957).
Within New York City and on Long Island, Pileated Woodpeckers were breeding
residents until the early to mid-1800s (Buckley et al. 2018). They disappeared from
the region as a whole following its rapid urbanization during the mid-1800s, and
there were no published sightings between 1879 and 1934 (Buckley et al. 2018, Bull
1964). Although they were not known to breed in the area during the 20th century
(Buckley et al. 2018, Bull 1964, Cruikshank 1942, Griscom 1923), as areas just
to the north of the region regained their forest cover, Pileated Woodpeckers began
to occur in New York City and on Long Island with some regularity as non-breeding
visitors during the spring and fall (Buckley et al. 2018). As Pileated Woodpecker
records increased, Bull (1964), documented a concurrent increase in the number
of breeding Pileated Woodpeckers in Westchester County, NY, and counties across
the Hudson River in New Jersey. By the 1960s, they were breeding just outside of
Bronx County, NY, at Grassy Sprain Reservoir in Westchester County (~8 km from
Bronx County; Buckley et al. 2018).
Pileated Woodpeckers were reported from Staten Island, NY, several times dur-
ing the 20th century as non-breeding visitors. In late-April and throughout May
1989, however, a Pileated Woodpecker was found occupying a territory in the High
Rock Park section of the Greenbelt, a large old-growth greenspace in central Staten
Island. On 31 May 1989, a single egg was found on the ground in front of the cavity,
after the Pileated Woodpecker was observed being mobbed by a pair of Colaptes
auratus auratus L. (Yellow–Shafted Northern Flicker). This egg was recovered and
placed in the collection of the Staten Island Museum by E.W. Johnson and R.V.
Matarazzo (Accession number B1989.5).
Here, we detail the re-establishment of Pileated Woodpeckers in New York City,
taking place in 2 old-growth greenspaces on Staten Island from 2015 to 2019. First,
we use linear discriminant analysis to verify that the egg found in High Rock Park
in May 1989 was laid by a Pileated Woodpecker, demonstrating that this species

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was likely attempting to breed in the region and going unobserved or unreported.
Second, we discuss their status in Long Island counties (Nassau and Suffolk) adja-
cent to New York City, where Pileated Woodpeckers are now occurring with some
regularity. The presence of this species in the most populous city and region in the
United States highlights the importance of unfragmented greenspaces for the pres-
ervation of biodiversity in urban areas.
Field–site Description
New York City is a densely populated metropolis composed of 5 counties,
with between about 500,000 and 2,500,000 residents each (United States Census
Bureau 2019). Prior to the widespread urbanization that took place during the
19th and 20th century, New York City had a mix of forest, grassland, and marsh
habitats, each of which were expansive and hosted a diverse array of species
(Sanderson and Brown 2007). Centuries of urbanization left each county with
varying percentages of remaining tree canopy and tree density (per acre; O’Neil-
Dunne 2012). Staten Island, Richmond County, the southernmost and least
populated county in New York City, has both the highest percent of tree canopy
cover remaining (30%) and the highest density of trees per acre (67.9 trees/acre)
(O’Neil–Dunne 2012).
We monitored Pileated Woodpeckers within 2 greenspaces on Staten Island: the
Greenbelt (Fig. 1c), and North Mount Loretto State Forest (Fig. 1b). In 1984, New
York City formally established the Greenbelt, a ~1214-ha (~3000-acre) greenspace
that spans the center of Staten Island (New York City Department of Parks and Rec-
reation 2019). The dominant tree species in the Greenbelt include Carya tomentosa
(Lam. Ex Poir) (Mockernut Hickory), Quercus rubra L. (Red Oak), Acer rubrum
L. (Red Maple), and Liquidambar styraciua L. (American Sweetgum). North
Mount Loretto State Forest is a 30-ha (74-acre) mixed hardwood forest on the south
shore of Staten Island (New York State Department of Environmental Conservation
2019). Dominant tree species include Acer saccharinum L. (Silver Maple), Fraxi-
nus pennsylvanica (Marshall) (Green Ash), Ulmus americana L. (American Elm),
and Liquidambar styraciua L. (American Sweetgum).
Methods
We measured the length (mm) and width (mm) of Egg B1989.5 in the Staten
Island Museum oology collection using Vernier calipers. The egg measured 32.50
mm x 24.50 mm. To determine the probability that the egg belonged to a Pileated
Woodpecker, we conducted a linear discriminant analysis (LDA) between eggs
of Yellow–shafted Northern Flicker—a widespread species in the eastern United
States with similar egg size and nesting ecology—and eggs of Pileated Wood-
peckers using the package ‘MASS’ (Venables and Ripley 2002) in the R statistical
programming language (version 4.0.2; R Core Team 2019). LDA is useful in
identifying species when there is overlap in their distinguishing characteristics
(see Sneath and Sokal 1973). For our comparison, we also measured Pileated

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Woodpecker (n = 93) and Yellow–shafted Northern Flicker eggs (n = 97) from
the oology collection at the American Museum of Natural History using the same
methods and calipers above.
Between 2014 and 2017, we made periodic observations of Pileated Woodpeck-
ers within the Greenbelt and North Mount Loretto State Forest. Upon the discovery
of a nest within the Greenbelt in 2018, we made weekly visits to the site during
the breeding season. To document the changing status of the species in the region,
we compiled records from the eBird citizen science database (Sullivan et al. 2009)
and from the New York State Ornithological Society’s peer–reviewed journal The
Kingbird. For historical data, we used a combination of The Kingbird seasonal re-
views of sightings and regional ornithological texts such as Buckley et al. (2018),
which includes an exhaustive list of non-breeding records for Bronx and New York
counties from 1939–2017.
Figure 1. (a) Map of Staten Island, New York; stars denote the 2 main study areas. (b) North
Mount Loretto State Forest, Staten Island. (c) The Greenbelt, Staten Island.

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Results
Verifying 1989 nesting attempt
The mean group length of Piliated Woodpecker eggs was 32.4 ± 0.01 mm and
the mean group width was 23.9 ± 0.01 mm. The mean group length of Yellow-
shafted Northern Flicker eggs was 28.4 mm and the mean group width was 21.8
mm. Egg length (t = -17.888, df = 147.94, P <0.001) and egg width (t = -20.129, df
= 147.03, P < 0.001) were signicantly different between species.
The results of the LDA indicated that Pileated Woodpecker eggs are signicant-
ly larger in length and width than Yellow-shafted Northern Flicker eggs. The LDA
explained 99.9% of the variation between groups, and predicted class membership
of Egg B1989.5 as belonging to Pileated Woodpecker (see Fig. 2). Cross validation
of the LDA model indicated that our LDA correctly predicts species classication
with 96.8% accuracy (P < 0.001).
Figure 2. Discrimination between Pileated Woodpecker egg measurements (n = 93) and
Yellow-shafted Northern Flicker egg measurements (n = 97).

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Successful establishment on Staten Island
Mount Loretto State Forest. On 23 August 2015, we observed and documented 2
hatch-year Pileated Woodpeckers associating with an adult bird within North Mount
Loretto State Forest (Fig. 3a). Following this sighting, we observed the hatch-year
Figure 3. (a) Two hatch-year Pileated Woodpeckers at North Mount Loretto State Forest,
Staten Island, with an arrow pointing to the bird obscured from view. (b) Pileated Wood-
pecker excavating a cavity within the High Rock section of the Greenbelt on Staten Island.

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birds repeatedly in the same location, last being sighted on 12 October 2015. Dur-
ing the following April, we again found a pair within the North Mount Loretto State
Forest and observed a single hatch-year bird in the same area on 24 August 2016.
In 2017, while we regularly observed Pileated Woodpeckers, we did not nd any
hatch-year birds nor any potential roosting or nesting cavities. In 2018 and 2019, we
observed a pair in the state forest, but no young were known to have edged.
The Greenbelt. On 8 April 2018, local bird watchers found a pair of Pileated
Woodpeckers excavating and later occupying a cavity within the Greenbelt (Fig.
3b). We did not monitor the nest regularly throughout the summer; however, we
observed at least one member of the pair returning to the nest site with food during
June. For the remainder of the breeding season, we did not observe any Pileated
Woodpeckers within the Greenbelt. On 9 September, 1 adult bird was found at High
Rock Park (C. Ricker, NYC Department of Parks and Recreation, NY, NY, pers.
comm.), but none were found for the remainder of 2018. On 26 March 2019, J.R.
Ramírez-Garofalo and S.R. Curley found that the original nest-site and cavity were
being occupied by Pileated Woodpeckers. On the morning of 27 May, we observed
the adult female foraging in the vicinity of the nesting site, and then feeding 1
hatch-year bird in the nest. On 2 July, we observed both adults and a hatch-year
bird in the same area. In 2018 and 2019, we made simultaneous observations of
pairs of Pileated Woodpecker within Mount Loretto State Forest and the Greenbelt,
indicating that there were in fact 2 pairs of birds involved in these sightings.
Current status elsewhere in New York City and Long Island
From 1935–2019, Pileated Woodpeckers mainly occurred in the large parks
within Bronx County and northern New York County, with 13 records for Van Cor-
tlandt Park, 6 for Bronx Park, and 5 for Inwood Hill Park (Table 1). They occurred
very rarely on Long Island, with 14 records for Queens, Nassau, and Suffolk coun-
ties combined, and none for Kings county. Although no Pileated Woodpeckers were
Table 1. Non-breeding records of Pileated Woodpeckers in New York City and Long Island from
January 1935 through February 2019. Excludes Staten Island records after 2014 when they became
permanent residents and evidently breeding. [Table continued on following page.]
Date Locality County Source
Late January 1935 Woodrow Richmond Davis 1937
20 April 1939 Bronx Park Bronx Bull 1964
21 April 1940 Bronx Park Bronx Buckley et al. 2018
22 June 1947 Sag Harbor Suffolk Bull 1964
1951 Staten Island Richmond SI Museum Archives
12 April 1959 Fort Tryon Park New York Bull 1964
23 April 1961 Inwood Hill Park New York Bull 1964
9 April 1962 Van Cortlandt Park Bronx Bull 1964
24 April 1964 Bronx Park Bronx Buckley et al. 2018
17 May 1971 Fieldstone Bronx eBird
4 May 1978 Bronx Park Bronx eBird
11 December 1981– Forest Hill Park Queens Levine 1998
1 May 1982

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Table 1, continued.
Date Locality County Source
April 1984 Van Cortlandt Park Bronx Buckley et al. 2018
March – June 1985 Van Cortlandt Park Bronx Buckley et al. 2018
April 1986 – April 1989 Pelham Bay Park Bronx Buckley et al. 2018
6 May 1986 Fisher’s Island Suffolk eBird
13 August 1987 Fisher’s Island Suffolk eBird
April –May 1989 High Rock Park Richmond This study
18 September 1990 Van Cortlandt Park Bronx Buckley et al. 2018
18 September 1994 Van Cortlandt Park Bronx Buckley et al. 2018
31 March 1995 Van Cortlandt Park Bronx Buckley et al. 2018
8 May 1996 Fisher’s Island Suffolk eBird
1 November 1997 Fisher’s Island Suffolk eBird
13 May 1998 Van Cortlandt Park Bronx Buckley et al. 2018
9 April 1999 Van Cortlandt Park Bronx Buckley et al. 2018
28 April 1999 Riverdale Bronx Buckley et al. 2018
28 April 1999 Clove Lakes Park Richmond Schiff and Wollin 1999
20 April 2000 High Rock Park Richmond Schiff and Wollin 2000
29 April 2004 Inwood Hill Park New York eBird
15 October 2006 Brookhaven National Suffolk Lindsay and Mitra 2007
Laboratory
24 February 2007 Buck's Hallow Richmond eBird
18 April 2009 Edenwald Bronx eBird
April 2011 Van Cortlandt Park Bronx Buckley et al. 2018
31 December 2011– Matinecock Nassau eBird
7 January 2012
7 March 2012 Buck’s Hallow Richmond eBird
5 May–15 September 2012 Bronx Park Bronx Buckley et al. 2018
19 –21 April 2013 Inwood Hill Park New York eBird
4 September 2013 Van Cortlandt Park Bronx Buckley et al. 2018
3–13 March 2014 North Mount Loretto State Richmond eBird
Forest
3 April 2014 Van Cortlandt Park Bronx Buckley et al. 2018
9 April 2014 Fort Tryon Park New York eBird
12 April 2014 Van Cortlandt Park Bronx eBird
13 April 2014 Inwood Hill Park New York eBird
4 May 2014 Oakland Lake Queens Ausubel 2014
5 May 2014 Valley Stream State Park Nassau Ausubel 2014
31 October 2014 North Mount Loretto State Richmond eBird
Forest
5 April 2015 Bethpage Nassau Ausubel 2015
27 June 2015 Bronx Park Bronx Buckley et al. 2018
18 October 2015 Pelham Bay Park Bronx Buckley et al. 2018
16 April 2016 Wave Hill Bronx eBird
25 March 2017 Inwood Hill Park New York eBird
4 –24 February 2018 Melville (private residence) Suffolk eBird
7 April 2018 Pelham Bay Park Bronx eBird
25 May 2018 Caumsett State Park Suffolk eBird
5 December 2018– Area around Pelham Bay Park Bronx eBird
4 April 2019
10 February 2019 Shu Swamp Nature Preserve Nassau eBird
8 September 2019 Van Cortlandt Park Bronx eBird

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found breeding outside of Staten Island, several individual birds lingered in some
of the larger parks (e.g., Pelham Bay Park) well into the breeding season (Table 1,
Fig. 4).
Discussion
The results of our LDA indicate that Pileated Woodpeckers laid 1 egg but did
not edge any young on Staten Island in 1989. This initial breeding attempt, which
was previously unpublished, occurred at a time when Pileated Woodpeckers were
either far less numerous (New Jersey) or completely absent (all of Long Island and
New York County) in the surrounding areas. This nding suggests that Pileated
Woodpeckers may have been trying to establish in some forested areas on Staten
Island and elsewhere in New York City and on Long Island, but were going un-
der- or undetected. Although Pileated Woodpeckers are large and noisy, there was
limited observer coverage on Staten Island, as well as in Bronx County, where there
is suitable habitat (Buckley et al. 2018). Community science applications like eBird
(https://ebird.org/home) and iNaturalist (https://www.iNaturalist.org) are rapidly
contributing to the effort to document the shifting distribution of species and will
aid in documenting further expansion by Pileated Woodpeckers in New York City
and Long Island. They will also be key to determining if Pileated Woodpeckers will
continue to establish on Staten Island.
Concurrent to the increases in their northeastern North American population,
Pileated Woodpeckers expanded their range north and eastward. This expansion
is shown by Canadian breeding bird atlas data for southern Québec (Robert et
al. 2019), Ontario (Cadman et al. 2007), and the Maritime Provinces (Stewart et al.
2015). Similarly in Connecticut, there has been an apparent eastward expansion
Figure 4. Map of New York City and Long Island. Black triangles indicate parks or greens-
paces where Pileated Woodpeckers have occurred at least once from 1935 to 2019.

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since the state’s rst breeding bird atlas survey (C. Elphick, University of Con-
necticut, Storrs, CT, pers. comm.). In Vermont (Renfrew 2013) and New York
(McGowan and Corwin 2008), data indicate an increase in block occupancy be-
tween each states’ rst and second breeding bird atlas surveys. In New Hampshire
(Keith and Fox 2015) and Massachusetts (Kamm et al. 2013), Pileated Woodpeck-
ers have become increasingly common in both suburban and urban areas, as well as
along the coast.
Pileated Woodpeckers are often depicted as being resident species, yet the pat-
tern of prospecting we describe suggests substantial dispersal is taking place. The
re-colonization of their pre-industrial range, and colonization of new regions like
the Canadian Maritimes, is likely driven by the exploratory behavior and dispersal
of (probably young) birds. For example, Peterjohn (1989) noted the autumn and
spring occurrence of this species at so–called “migrant traps” along Lake Erie,
the timing of which is suggestive of breeding territory prospecting (see Reed et al.
1999). Further evidence of their dispersal ability is displayed by vagrant records
on 22 September 1925 and 26 December 1954 from Block Island, RI, and Nan-
tucket Island, MA, respectively (Ferren and Veit, in press; Veit and Petersen 1993),
both of which are located over 10 km from the mainland. Recent records from the
Sandy Hook and Cape May peninsulas in New Jersey (eBird data) represent similar
dispersal over bodies of water, or large tracts of habitat not typically suitable for
Pileated Woodpeckers such as barrier beaches. Long-distance dispersal, which of-
ten drives the overall rate of expansion (Kot et al. 1996, Veit and Lewis 1996), was
likely a major factor in the re-establishment of Pileated Woodpeckers throughout
northeastern North America, as opposed to only short-distance movements, par-
ticularly in the densely urbanized coastal regions where forest fragments may be
the only suitable habitat for tens of kilometers.
The presence of breeding Pileated Woodpeckers in New York City calls atten-
tion to the importance of large, unfragmented greenspaces within urban areas.
Greenspaces like Staten Island’s Greenbelt may be of importance to conserva-
tion of species expanding their range in a rapidly urbanizing landscape. The
presence of greenspaces can increase local biodiversity (Aronson et al. 2017),
provide habitat for area-sensitive species (Morrison and Chapman 2005, Mört-
berg and Wallentinus 2000, Park and Lee 2000), and act as dispersal corridors
through urban areas (Bolgerm et al. 2001). Staten Island, while having the high-
est percentage of tree canopy cover in New York City (O’Neil-Dunne 2012), has
been rapidly and widely developed since the 1960s (Siebenheller 1981). Develop-
ment has continued to encroach on areas within the Greenbelt, to the detriment of
forest-interior specialists (Dowd 1992). Park and greenspace management plans
should consider the presence of Pileated Woodpeckers as an indicator of good for-
est health (McClelland and McClelland 1999) and as general indicators of forest
bird diversity (Mikusiński et al. 2001).
Pileated Woodpeckers have received little study in northeastern North America
(Flemming et al. 1999). The Pileated Woodpeckers breeding on Staten Island pres-
ent a unique opportunity to study their nesting ecology in the region and to guage

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the needs of a forest specialist in an urban setting. We could enhance such studies
using color-banding and radio telemetry, which would allow study of their dispersal
behavior in the region (see Tomasevic and Marzluff 2018). Such research would,
in turn, help determine whether Staten Island-born Pileated Woodpeckers are dis-
persing into other areas in New York City and Long Island, or if they are recruiting
into the New Jersey population. Community scientist participation would supple-
ment re-sighting efforts and could also be valuable for collecting demographic data
through nest-watching projects (Bonney et al. 2009, Evans et al. 2005).
Acknowledgments
We thank 2 anonymous reviewers for comments that greatly improved the quality of
this manuscript. We further thank Colleen Evans of the Staten Island Museum for facilitat-
ing our visits to the oology collection, and Paul Sweet of the American Museum of Natural
History for facilitating our visit to compare and measure eggs in the Museum’s collection.
We thank Thomas Dulski and Isaac Grant for alerting us to the presence of nesting Pileated
Woodpeckers in Staten Island’s Greenbelt. P.A. Buckley, Michael Shanley, Seth Wollney,
Clifford Hagen, and James Rossi also provided information on unpublished sightings of
Pileated Woodpeckers. Zachary Johnson assisted us in nding relevant historical informa-
tion from local natural history notes and publications. Regular information on activity by
Pileated Woodpeckers within Staten Island’s Greenbelt was provided to us by Christopher
Ricker and Karen Roos of the New York City Department of Parks and Recreation.
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