Breeding Bird Patterns and Species Turnover within the Central Business District of a Town: Results from Rockingham, North Carolina, USA
Douglas B. McNair1*
135 Rowell Road, Wellfleet, MA, USA 02667. *Corresponding author.
Urban Naturalist, No. 44 (2021)
Small cities or towns, and in particular central business districts in southeastern North America, are still underrepresented in breeding bird surveys. Using the spot-mapping method, I documented 21 species (17 native suburban adapters, 4 exotic urban invaders) nested within a 25-block study area (42.4 ha) of downtown Rockingham (population ~9,000) in south-central North Carolina in 1994. The 156 breeding pairs had a density of 147.1 territories (T)/40 ha (native: 90.5 T/40 ha, exotics: 56.6 T/40 ha). The 6 most abundant species were 3 exotics (European Starling: 20.8 T/40 ha; House Sparrow: 18.9 T/40 ha; House Finch: 12.3 T/40 ha) and 3 natives (Common Grackle: 17 T/40 ha; Mourning Dove: 14.6 T/40 ha; American Robin: 12.7 T/40 ha). I documented 433 nests and confirmed breeding records, 324 (75%) of which were open-cup nests. Each species nested entirely (or almost entirely) in vegetation or on or within buildings and other anthropogenic structures. Only House Finches, before an outbreak of mycoplasmal conjunctivitis reached Rockingham, nested at almost an equal number of vegetative and building sites. Median species richness and number of nest records were significantly greater in vegetation than at buildings; only 4 of 25 blocks (16%) had a higher proportion of their nest records at buildings. Supplemental surveys conducted 18–23 years later (2012, 2016–2017) that focused on species turnover documented the loss of 2 of the 3 most numerous native species (Common Grackle, American Robin), even though habitat remained largely unchanged.
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2021 No. 44
2021 Urban Naturalist 44:1–19
Breeding Bird Patterns and Species Turnover within the
Central Business District of a Town:
Results from Rockingham, North Carolina, USA
Douglas B. McNair1*
Abstract - Small cities or towns, and in particular central business districts in southeastern North
America, are still underrepresented in breeding bird surveys. Using the spot-mapping method, I documented
21 species (17 native suburban adapters, 4 exotic urban invaders) nested within a 25-block
study area (42.4 ha) of downtown Rockingham (population ~9,000) in south-central North Carolina
in 1994. The 156 breeding pairs had a density of 147.1 territories (T)/40 ha (native: 90.5 T/40 ha,
exotics: 56.6 T/40 ha). The 6 most abundant species were 3 exotics (European Starling: 20.8 T/40
ha; House Sparrow: 18.9 T/40 ha; House Finch: 12.3 T/40 ha) and 3 natives (Common Grackle: 17
T/40 ha; Mourning Dove: 14.6 T/40 ha; American Robin: 12.7 T/40 ha). I documented 433 nests and
confirmed breeding records, 324 (75%) of which were open-cup nests. Each species nested entirely
(or almost entirely) in vegetation or on or within buildings and other anthropogenic structures. Only
House Finches, before an outbreak of mycoplasmal conjunctivitis reached Rockingham, nested at
almost an equal number of vegetative and building sites. Median species richness and number of nest
records were significantly greater in vegetation than at buildings; only 4 of 25 blocks (16%) had a
higher proportion of their nest records at buildings. Supplemental surveys conducted 18–23 years later
(2012, 2016–2017) that focused on species turnover documented the loss of 2 of the 3 most numerous
native species (Common Grackle, American Robin), even though habitat remained largely unchanged.
Urbanization connotes changes in landscape (or environment) caused by urban development
(Marzluff 2001, Marzluff et al. 2001). Biotic homogenization is characteristic of urban
landscapes, where resident habitat generalist avian species thrive and Neotropical and Palearctic
migrants sharply drop in richness (Aldrich and Coffin 1980, Aronson et al. 2014, Beninde
et al. 2015, Blair 2004, Chace and Walsh 2006, Clucas and Marzluff 2015, Emlen 1974,
Kelcey and Rheinwald 2005, La Sorte and McKinney 2007, Leston and Rodewald 2006).
Most avian studies of biotic homogenization in urban landscapes of North America have been
conducted in residential areas, parks, riparian corridors, and forest fragments of suburbia (Aronson
et al. 2014, Belaire et al. 2015, Beninde et al. 2015, Chace and Walsh 2006, Clucas and
Marzluff 2015, Duren et al. 2017, Leston and Rodewald 2006, Marzluff and Rodewald 2008,
Schneider and Miller 2014), including locations in the southeastern United States (Aldrich
and Coffin 1980, Rohwer and Woolfenden 1969, Thorington and Brand 2014, Woolfenden
and Rohwer 1969). For example, Woolfenden and Rohwer (1969) emphasized relationships
between the density of breeding pairs of avian species and a variety of landscape variables
within the urbanized matrix of Tampa, Florida. More recently, in Winston-Salem, North Carolina,
breeding bird censuses (hereafter, BBCs) using spot-mapping of territories over 2 years
documented 60 breeding species in a large park (77-ha) surrounded by suburban residential
housing (Thorington and Brand 2014). In southeastern North America, no BBCs, other than
in large urban areas, have been conducted in a town and none in a central business district.
The closest locations known to the author have been towns in eastern Kansas (Baldwin City)
and northern Ohio (Carey), where both residential and commercial areas were censused (Cink
135 Rowell Road, Wellfleet, MA, USA 02667. *Corresponding author: email@example.com.
Manuscript Editor: Michael Strohbach
2021 No. 44
1983, Claugus 1983). Overseas, in the Czech Republic (Kučera et al. 2015) and in Germany
(Nicolai and Wadewitz 2003), a few recent assessments of avian communities that included
central business districts have been published for towns similar in size or somewhat larger
compared to Rockingham, Richmond County, North Carolina.
Rockingham has long been a bird-friendly town, with posted signs announcing it was
designated a bird sanctuary in the early 1950s (Barber 1952). Rockingham, unlike most large
urban areas of North Carolina, but like most towns and small cities in the Pee Dee region of
the Carolinas, has a stagnant economy, is not expanding geographically, and has not increased
in population for over 35 years (Hutchinson et al. n.d.; US Census Bureau 2020). The landscape
of Rockingham’s central business district and outlying urban areas has remained largely
unchanged, unlike larger urban centers with dynamic economies and expanding populations
where most avian studies on urbanization effects have been conducted (e.g., Seattle, Washington;
Clucas and Marzluff 2015). Thus, birds in Rockingham have had low urbanization pressure,
but they have faced pressure from other factors. For example, the spread of mycoplasmal
conjunctivitis in Haemorhous mexicanus (P.L. Statius Müller) (House Finch) populations, an
urban invader, began in eastern North America in 1994 (Badyaev et al. 2012). Furthermore,
several species of suburban adapters that may breed in towns have had regional range expansions
(McNair 2019) or contractions (McNair 2015) that have included Rockingham over the
past decade. The scarcity of studies from towns in rural North America, stagnant development,
and dynamics of other factors create a need for a better understanding of avian species relationships
in smaller urban landscapes such as Rockingham. The main objectives of this study
were to (1) examine patterns of functional guilds, bird species richness, and nest densities in
relationship to built-up land and vegetation and (2) discuss species turnover 2 decades after
first sampling at Rockingham in 1994.
Study Area Description
Rockingham, North Carolina, the county seat of Richmond County, is a micropolitan
statistical area. Rockingham was founded in 1774 at Cole’s Hill (Hutchinson 1998), a bluff
rising between Hitchcock and Falling Creeks, along the contact zone (Fall Line) where the
Piedmont meets the Sandhills, a subregion of the Coastal Plain. The study area is centered
on the ancient location of Cole’s Hill, which comprises the central business district of
Rockingham (although most commercial development in the micropolitan statistical area
now occurs along US BUS 74 between Rockingham and Hamlet, 8 km east-southeast). The
estimated population of Rockingham, the county’s most populous community, was 8994 in
2018, a decline of 4.3% from its population of 9399 in 1990 (US Census Bureau 2020).
The study area (42.4 ha; Fig. 1) on Cole’s Hill in the central business district of Rockingham
(henceforth called downtown Rockingham: Latitude 34.939528 N, Longitude
79.761236 W; 87 m asl) excludes town blocks of adjacent residential areas. The study
area, on flat to moderately steep terrain, is bounded on the west and south sides by the
intersection of US 220 and US BUS 74. These 2 major highways connect to town streets
that define the northern and eastern boundaries of the study area (Fig. 1). The southeastern
boundary of the study area that encloses business district parcels includes a ruderal habitat
patch dominated by Pueraria (Kudzu). The closest points along Hitchcock and Falling
Creeks are located 60 m and 40 m, respectively, outside the northwestern and southeastern
boundaries of the study area.
The study area does not contain commercial buildings taller than 3 stories, riparian or
other aquatic habitats, forests, woodlands, farmland, parks, cemeteries, allotment gardens,
2021 No. 44
or industrial areas. Downtown Rockingham contains a mixture of cropped (maintained)
remnant native vegetation and exotic vascular plants. Plantings on properties in the central
business district usually adhered to recommendations of regulations by the City of Rockingham
(2020). The tallest tree within the study area was 30 m high.
I digitized individual block size within the 25-block study area from recent (2016) high
resolution aerial imagery, which I analyzed in ESRI’s ArcGIS 10.4 software. The mean and
median block sizes were 1.70 ha (± 0.20 SE) and 1.46 ha (interquartile range: 1.20–2.07
ha), respectively (min–max range: 0.18–4.12 ha). I used street centerlines to demarcate the
boundaries between blocks except along the eastern boundary where vegetation formed the
outer boundary of 3 blocks (nos. 4–5, 16).
Land Cover Data
The City of Rockingham (J.E. Armstrong, past Planning Director, City of Rockingham,
NC, pers. comm.), in 1994, provided the area of buildings within each block, which totaled
6.59 ha (15.5% of the total study area); this amount does not include streets, pavements, and
Figure 1. The 25-block study area in downtown Rockingham, Richmond County, North Carolina, with
each block represented by a number, largely consists of buildings and other sealed surfaces, with smaller
areas of a mixture of native and exotic vegetation, including lawns. The 0.4-km perimeter beyond
the study area boundary is designated by a black dashed line. The source of the image is from the North
Carolina Orthoimagery Program (date of imagery 8 March 2015) and is being used with permission.
2021 No. 44
other impervious surfaces on the ground. The mean and median areas of buildings within
blocks were 0.26 ha (± 0.036 SE) and 0.24 ha (interquartile range: 0.12–0.37 ha), respectively
(min–max range: 0.02–0.66 ha).
I developed a vegetation index for the diverse mixture of remnant native vegetation and
landscaped plantings by overlaying an acetate grid of mylar squares on magnified aerial
photographs of each of the 25 blocks at a scale of 1:840 (2.54 cm:21.336 m; original measurements:
1 in:70 ft). This vegetation index relied upon estimation of the spatial coverage
of vascular plants within each block. I excluded measurements of any exaggerated images
of vascular plants. If vegetation filled at least 50% of a mylar square, I scored it as 1 square.
This index excluded plants less than 1 m tall and ground vegetation, including lawns, where
birds did not nest.
I reassessed the study area 18 to 23 years later (2012, 2016–2017) for any changes in
habitat and landscape characteristics (including buildings and paved surfaces) that may
have occurred since 1994. I examined photographs and field notes I took in 1994 to compare
with field inspections during the 3 later years. I also compared individual blocks from past
(1993) and recent (2013, 2017) aerial imagery for all 25 blocks and used the vegetation
index as I did in 1994 to quantify vegetative changes.
Species nomenclature for scientific names, authorities, and common names follows
Chesser et al. (2019) and ITIS (2020). I conducted avian surveys on 94 days from 27 March
to 18 September 1994; field effort was concentrated from April through July (236.75 of
252.25 h; 94%). Less search effort occurred before noon (79.5 h; 31.5% of total) than
afterward. The BBC uses the spot- or territory mapping method to estimate densities (Aldrich
and Coffin 1980, Gardali and Lowe 2006, Leston and Rodewald 2006, Thorington
and Brand 2014). Following this protocol during the peak breeding period (April to early
June) for species that maintained exclusive multi-purpose territories, I rounded the number
of territories to the nearest half territory. There were 8 species that may not maintain exclusive
multi-purpose territories but may only defend nesting-station territories as isolated
pairs (although several species may nest semi-colonially): Columba livia var. domestica
J.F. Gmelin (Feral Pigeon), Zenaida macroura (L.) (Mourning Dove), Chaetura pelagica
(L.) (Chimney Swift), Cyanocitta cristata (L.) (Blue Jay), Sturnus vulgaris (L.) (European
Starling), Passer domesticus (L.) (House Sparrow), House Finch, and Quiscalus quiscula
(L.) (Common Grackle). In addition to spot-mapping, I used the location of active nests or
confirmed breeding attempts without direct evidence but based on observations of behavior
(e.g., adults of cavity-nesting species repeatedly entering a cavity with food) to confirm
the number of territories for these 8 species. The number of pairs for Feral Pigeons was
considered to be half of their individual numbers recorded in 1 count (Luniak 1994). For
Chimney Swifts, the number of chimneys that were regularly used constituted the number
of breeding pairs since only 1 pair nests per chimney (Steeves et al. 2014). I standardized
breeding densities within the study area by calculating the number of territories per 40 ha;
this calculation included species with less than 3.0 territories.
I also conducted intensive nest searches to complement use of the spot-mapping method
to delineate territories. Nest records had the advantage that they could be consigned to an
individual block and its specific vegetative and anthropogenic characteristics unlike many
territories, which encompassed more than 1 block. Most effort was devoted to obtaining information
on open-cup breeding species (which included 1 urban invader, the House Finch)
rather than cavity-nesting species. The latter’s nests were generally inaccessible, although
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their breeding activity was closely monitored. The 12 most numerous species (see Results)
were multiple-brooded, except for the Common Grackle, which is usually single-brooded
(Peer and Bollinger 1997).
I supplemented 1994 data by obtaining additional information in 2012 and 2016–2017. I
conducted these supplemental surveys on 24 days from 7 April to 2 June for a total of 35.5
h. Search effort before (17 h; 48%) and after noon (18.5 h; 52%) was approximately equal.
The focus was on documentation of species turnover (presence or absence), the number of
territories of species that were scarce in 1994, and the number of territories of new species
discovered during the later period. Otherwise, I did not map territories or attempt to find
nests of the more numerous species. These efforts allowed me to document any species
turnover that occurred between 1994 and the 3 later years.
During avian surveys from March to September 1994, I obtained weather data at Hamlet,
North Carolina (Latitude 34.8872 N, Longitude 79.6922 W, 107 m asl) because comparable
weather data were not available from Rockingham (NRCC 2017). In comparison to Normals
period (1981–2010) data during these 7 months, temperature and precipitation in 1994 were
slightly cooler and wetter than average. The exception was rainfall of 38.05 cm in July 1994,
the highest rainfall during July over this 30-year period.
Functional guilds. Breeding birds in the central business district of Rockingham can be
loosely categorized into 2 functional guilds based on their differing responses to urbanization—
urban invaders and suburban adapters (Marzluff and Rodewald 2008; adopted and
developed from Blair 1996). Urban invaders (e.g., European Starling) are rare in natural
areas and abundant and fecund in urban areas. Suburban adapters (e.g., American Robin L.
[Turdus migratorius]) are typically adapted to diverse, young, edge, and disturbed habitats
(Marzluff and Rodewald 2008). These species would include the Chimney Swift, which
readily occupies buildings in suburban and urban environments.
Biodiversity measures. I calculated species richness (number of different species present),
which is the most studied attribute of urban communities (Marzluff and Rodewald
2008), densities of breeding birds, and number of nests and confirmed breeding attempts
(henceforth called nest records) for the entire 25-block study area and, where applicable, for
each individual block in downtown Rockingham in 1994. The number and location of nest
records were used to measure each species distribution in the study area. I examined scatterplots
and conducted non-parametric Spearman’s correlation tests to assess monotonic links
between these 3 biodiversity measures (species richness, breeding density, number of nest
records). Species richness among all blocks was strongly correlated with the total number
of nest records, the number of nest records in vegetation, and the number of nest records at
buildings (rS = 0.87, 0.83, and 0.90, respectively; all P < 0.001). Furthermore, the breeding
bird densities of all species was strongly correlated with the number of nest records (rS =
0.83, P < 0.001). Thus, use of the number of nest records in subsequent analyses was justified.
Effects of buildings and vegetation. Landscape regulations of the City of Rockingham
(2020) stipulate that 5 blocks (nos. 1, 2, 8, 16, 17; see Fig. 1) and a portion of properties
within 3 other blocks (nos. 9, 11, 21) should consist of a minimum of 12% landscaped
areas, which includes lawns. The remaining 17 blocks are exempted from this regulation,
and their landscaped areas can be less than 12%. Excluding lawns, the median difference
in landscaped areas between these 2 groups (≥12%, <12%), using the vegetation index,
was not significantly different (Mann–Whitney U = 37, 2-tailed test, P = 0.08). Therefore,
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I made no distinction in the proportion of spatial coverage of vascular plants in landscaped
areas among the 25 blocks.
I used 2-tailed non-parametric Wilcoxon signed-rank tests to examine median differences
between non-independent matched-pair observations (buildings, vegetation) from each
block, across all 25 blocks, for species richness and the number of nest records (I omitted
Block 1 for nest records because search effort in vegetation was inadequate). I examined
scatterplots and conducted non-parametric Spearman’s correlation tests to assess whether
building area and the vegetation index were monotonically linked to species richness or the
number of nest records across all blocks (omitting Block 1 for testing with the vegetation
index) for each of the 10 most abundant species.
I also conducted a first-order partial correlation analysis to examine the effects of building
area and the vegetation index on species richness and the number of nest records for all
species across all blocks, after adjustment for the effect of block size. I converted values
using a log10 (n + 1) transformation for 3 of 4 variables (species richness, nest records,
block size) in the building analysis and 2 of 4 variables (vegetation index, block size) in the
vegetation analysis so the data became normally distributed or more closely approximated a
normal distribution; data transformation eliminated outlier values and ensured homogeneity
of variances. All analyses were performed with statistical software available from McDonald
(2014) and Real Statistics Using Excel (2019).
Species richness. Twenty-one avian species (16 passerines) nested within the study area
of downtown Rockingham in 1994 (Table 1), including 13 open-cup species (10 residents, 3
Neotropical migrants) and 8 cavity species (6 residents, 2 Neotropical migrants). Seventeen
species were native suburban adapters, whereas 4 species were exotic urban invaders. An
additional 30 avian species visited the study area during the breeding season in 1994 but
did not breed (Appendix 1). Twenty-one of these 30 visitors plus an additional 20 species
bred within 0.4 km of the perimeter of the study area in 1994 (Appendix 1). These species
included Molothrus ater (Boddaert) (Brown-headed Cowbird), which was not detected
within the study area after 27 March.
Breeding densities. Breeding bird density for all 21 species during the peak season
(April–early June) was 156 territories (Table 1), a density of 147.1 territories (T)/40 ha
(native: 90.5 T/40 ha, 61.5%; exotics: 56.6 T/40 ha, 38.5%). Resident species accounted
for 146 territories (93.6%; 137.7 T/40 ha), whereas the 5 Neotropical migrants accounted
for 10 territories (6.4%; 9.4 T/40 ha), including 6 territories of Dumetella carolinensis
(L.) (Gray Catbird). The 6 most abundant species were 3 exotics (European Starling: 20.8
T/40 ha; House Sparrow: 18.9 T/40 ha; House Finch: 12.3 T/40 ha), which included 2
cavity species, and 3 natives (Common Grackle: 17 T/40 ha; Mourning Dove: 14.6 T/40
ha; American Robin: 12.7 T/40 ha). The Common Grackles nested as isolated pairs and
semi-colonially (in Block 25). Nine of the 21 breeding species (43%) each had only a
Nest records. I found 433 nest records in 1994 (Table 1), 324 (75%) of which were opencup
nests. These findings included nest records for all 7 native species present on single territories
in 1994. Only Thryothorus ludovicianus (Latham) (Carolina Wren) nest records in
1994 were proportionally under-recorded compared to their number of breeding territories
2021 No. 44
Distribution. The most abundant species were generally the most widely distributed
species within the 25-block study area, based on the total number of nest records of each
species among all blocks (Table 1; Fig. 2A: rS = 0.97, P < 0.001). Sixteen species breeding
in vegetative nest sites separated themselves into 3 groups: 8 sparsely distributed species
(including European Starling), 5 moderately distributed species (Gray Catbird, Toxostoma
rufum [L.] [Brown Thrasher], House Finch, Common Grackle, Cardinalis cardinalis [L.]
[Northern Cardinal]), and 3 widely distributed species (Mourning Dove, American Robin,
Mimus polyglottos [L.] [Northern Mockingbird]; Fig. 2B: rS = 0.99, P < 0.001). Nine species
breeding at buildings separated themselves into 3 groups: 4 sparsely distributed species
(including Mourning Dove and American Robin), 2 moderately distributed species (Feral
Pigeon, Carolina Wren), and 3 widely distributed species (European Starling, House Sparrow,
Table 1. Number of territories, territories per 40 ha, nest records, and city blocks with nests for 21 avian
species that bred within the 25-block study area in downtown Rockingham, North Carolina, in 1994.
Open-cup and open-air1 breeding species
Territories Territories/40 ha Nest records City blocks with nests
Zenaida macroura (L.)
15.5 14.6 56 19
Charadrius vociferus L.
1 0.9 2 1
Lanius ludovicianus L.
1 0.9 1 1
Cyanocitta cristata (L.)
1 0.9 1 1
Turdus migratorius L.
13.5 12.7 63 22
Dumetella carolinensis (L.)
6 5.7 31 11
Toxostoma rufum (L.)
9 8.5 26 12
Mimus polyglottos (L.)
11 10.4 51 21
Haemorhous mexicanus (P.L.
13 12.3 40 16
Icterus spurius (L.)
1 0.9 1 1
Quiscalus quiscula (L.)
18 17.0 25 10
Cardinalis cardinalis (L.)
6 5.7 26 14
Passerina caerulea (L.)
1 0.9 1 1
Subtotal 97 91.5 324 na2
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and House Finch; Fig. 2C: rS = 0.92, P < 0.001). The distributions of European Starlings and
House Sparrows at buildings were truncated at 12 blocks, despite their greater abundances
in comparison to the House Finch. European Starlings and House Sparrows were less widely
distributed compared to the 4 most widely distributed open-cup nesting species (including the
House Finch, which nested in a total of 16 blocks, vegetative and building sites combined).
Effects of Buildings and Vegetation
Median species richness and the number of nest records of matched-pair observations at
blocks, across all 25 blocks, were significantly greater in vegetation (species richness = 6,
interquartile range = 4–7, min–max = 0–9; number of nest records = 13, interquartile range
= 5–17, min–max = 0–33) than at buildings (species richness = 2, interquartile range = 1–3,
Table 1. Continued.
Cavity and crevice breeding species3
Territories Territories/40 ha Nest records City blocks with nests
Columba livia var. domestica J.F.
5 4.7 12 7
Chaetura pelagica (L.)
1 0.9 1 1
Melanerpes carolinus (L.)
1 0.9 1 1
Myiarchus crinitus (L.)
(Great Crested Flycatcher)
1 0.9 1 1
Sitta pusilla Latham
1 0.9 1 1
8 7.6 7 6
Sturnus vulgaris L.
22 20.8 47 14
Passer domesticus (L.)
20 18.9 39 12
Subtotal 59 55.7 109 na2
TOTAL 156 147.1 433 na2
1 The Mourning Dove builds a simple platform nest; the Killdeer laid its eggs in a scrape on a rooftop.
2 na = not applicable.
3 The Feral Pigeon builds a simple platform nest on ledges located in the interior of buildings; the Chimney
Swift builds stick nests glued together by saliva inside open chimneys or in interior walls of abandoned
buildings; the Carolina Wren usually builds domed nests with side entrances in crannies or other secluded
places; the House Sparrow builds open-cup to globular-shaped nests in crevices in a variety of anthropogenic
2021 No. 44
min–max = 0–7; number of nest records = 3, interquartile range = 1–10, min–max = 0–15;
species richness: Wilcoxon signed-rank test: Z = 3.86, n = 25, P < 0.001; number of nest
records: Wilcoxon signed-rank test: Z = 2.94, n = 24, P = 0.003). Four of 24 blocks (17%)
had a higher proportion of their nest records at buildings, 1 block had an equal proportion
(4%), and 19 of 24 blocks (79%) had a higher proportion of their nest records in vegetation
Building area and the vegetation index were positively associated with the number of
nest records at buildings and in vegetation, respectively, for each of the 10 most abundant
species. These trends were significant at buildings for 2 urban invaders (European
Starling, House Sparrow; rS range = 0.54–0.61, n = 25, P range = 0.001–0.006) and 4
suburban adapters (Mourning Dove, Gray Catbird, Brown Thrasher, Northern Cardinal;
Figure 2. (A) The number of blocks with nest records by the total number of nest records for each of 21
avian species; (B) the number of blocks with nest records by the number of nest records in vegetation for
each of 16 avian species; and (C) the number of blocks with nest records by the number of nest records
at buildings for each of 9 avian species that bred in downtown Rockingham, North Carolina, in 1994.
2021 No. 44
rS range = 0.43–0.59, n = 24, P range = 0.002–0.03). The weakest associations were for
the House Finch, at buildings (rS = 0.09, n = 25, P = 0.66) and in vegetation (rS = 0.001,
n = 24, P = 0.997).
Building area was positively associated with species richness and the number of nest
records at buildings for all species but the trends were not significant after adjustment for
the effect of block size (species richness: rp = 0.32, n = 25, P = 0.12; nest records: rp =
0.36, n = 25, P = 0.09; first-order partial correlation tests). However, examination of scatterplots
revealed that species richness was disproportionately high in Block 12 and disproportionately
low in Blocks 8 and 18, and there was a disproportionately high number
of nest records in Block 19. In contrast, the vegetation index was significantly associated
with species richness and the number of nest records in vegetation for all species, after
adjustment for the effect of block size (species richness: rp = 0.50, n = 24, P = 0.02; nest
records: rp = 0.49, n = 24, P = 0.02). Nonetheless, examination of scatterplots revealed a
disproportionately high number of nest records in Block 25 (where a Common Grackle
semi-colony was located) and a disproportionately low number in Block 16 (where the
Kudzu patch was located).
Land Cover Change
In 1994, the study area contained bird feeders at 4 locations in 4 blocks, bird baths at 3
locations in 3 blocks, and 1 nest box in 1 block. Isolated snags were present in ~5 blocks.
Figure 3. The proportion of the number of avian nest records at buildings for 24 blocks in downtown
Rockingham, North Carolina (Block 1 is omitted because search effort in vegetation was inadequate);
each block is denoted by a number from Figure 1. An equal proportion of nest records at buildings
and vegetation is indicated by a value of 0.5, a value >0.5 indicates a greater proportion at buildings,
and a value <0.5 indicates a greater proportion in vegetation.
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By 2012, 1 bird bath was still present in the study area, but bird feeders and nest boxes were
absent, and all isolated snags had been removed.
The sum total loss of vegetation in the later period compared to 1994, as measured by
the vegetation index, was 5.4%; thus, 94.6% of spatial coverage of vascular plants from
1994 was still present. Habitat and landscape characteristics in the 3 later years remained
unchanged since 1994 in 19 of 25 blocks (76%). Five of the 6 remaining blocks (nos. 5, 9,
11, 12, 21; see Fig. 1) had less vegetation and/or more buildings (including the first building
more than 3 stories high, the Richmond County Judicial Center in Block 9 where the
greatest vegetative loss occurred). Most loss of vegetation was removal of solitary trees
and shrubs and hedgerows, usually for expansion of parking lots. The sixth block (no. 16)
contained slightly more vegetation in the 3 later years than in 1994.
Species turnover in 1994 compared to the 3 later years was a net loss of 7 native species
(loss of 9, gain of 2). Territorial pairs of Common Grackles were absent from the study area
in 2012 and 2016–2017; I found 1 old nest (from 2011) in early April 2012 at the former
semi-colony in Block 25. Territorial pairs of American Robin were present in 2012 and
2016 but absent in 2017 (1 male sang on territory on 2 May, but that bird did not remain).
Seven species present as single territorial pairs in 1994 were absent during the 3 later years,
including 3 cavity nesting species (Melanerpes carolinus [L.] [Red-bellied Woodpecker],
Myiarchus crinitus [L.] [Great Crested Flycatcher], Sitta pusilla Latham [Brown-headed
Nuthatch]) and 4 open-cup nesting passerines (Lanius ludovicianus L. [Loggerhead Shrike],
Blue Jay, Icterus spurius [L.] [Orchard Oriole], Passerina caerulea [L.] [Blue Grosbeak]).
Two new native passerines (Stelgidopteryx serripennis [Audubon] [Northern Roughwinged
Swallow], Spizella passerina [Bechstein] [Chipping Sparrow]) nested within the
study area during the later years. Chipping Sparrows were present during all 3 years (3.0,
3.5, and 1.0 territorial pairs, respectively), and I discovered 2 nests in Quercus phellos L.
(Willow Oak) and Pinus taeda L. (Loblolly Pine). One pair of Northern Rough-winged
Swallows nested in a decommissioned long trailer de-hitched from a cab (Block 1) in 2016
and 2017; in 2017, the pair abandoned the site in late April when the trailer was re-commissioned
Two species (Chimney Swift, Charadrius vociferus L. [Killdeer]) that nested as single
pairs in 1994 also nested during the 3 later years. Three territorial pairs of Chimney Swifts
nested in 3 chimneys in 3 blocks in all 3 years. Killdeers, which had nested on rooftops in
downtown Rockingham since at least 1979, nested on the ground in 2017 (Block 11; adult
incubating 4 eggs on 15 May, though the nest failed shortly thereafter). Finally, ~2.0 territorial
pairs of Feral Pigeons were present in the later years in 2 blocks. Accipiter cooperii
(Bonaparte) (Cooper’s Hawk) preyed upon Feral Pigeons during the breeding season of 2011,
including pigeons that nested inside the Everett-McNair Store (Block 7; J.R. Massey, Jr., Assistant
City Manager, City of Rockingham, NC, pers. comm.). I observed a Cooper’s Hawk
preying on pigeons at a former hardware store (Block 1) during April–May 2017.
This study has extended the usefulness of BBCs (Engstrom and James 1984) by coupling
documentation of breeding territories by the spot-mapping method with the number of nest
records in urban blocks as part of the Rockingham BBC. Town blocks with the greatest areas
of buildings and the highest vegetation indices, respectively, were positively associated
2021 No. 44
with species richness and the number of nest records of species categorized as urban invaders
(Chace and Walsh 2006, Marzluff 2001) and suburban adapters (Beninde et al. 2015).
The probable underestimation of the distribution (based on the number of nest records)
of the Carolina Wren was not a critical shortcoming. Regardless, biotic homogenization
of avian species richness, composition, and breeding densities within the central business
district of Rockingham, North Carolina, is typical of the loss of biodiversity in most larger
urban landscapes worldwide (Marzluff 2001, Marzluff et al. 2001).
The most abundant birds on the Rockingham BBC in 1994 in blocks dominated by buildings
were 2 exotic resident cavity-nesting species (European Starling, House Sparrow).
This pattern is expected in central business districts of cities and towns, including those in
eastern North America (Aronson et al. 2014, Beninde et al. 2015, Chace and Walsh 2006,
Clucas and Marzluff 2015, Kelcey and Rheinwald 2005, Marzluff 2001). Nonetheless,
the number of blocks occupied by these 2 urban invaders at buildings was truncated at 12
blocks. Building condition, despite some vacancies, evidently set an upper limit to the number
of blocks European Starlings and House Sparrows could occupy. Variation in building
condition among blocks of different sizes is probably responsible for the lack of significant
associations between building area and species richness or the number of nest records.
Building condition in some blocks is still poor (e.g., Block 19, with a high number of nest
records), but construction of relatively new buildings (e.g., Block 8, with a low number of
nest records) and renovation and restoration efforts for over 20 years has slowly improved
building condition in downtown Rockingham (e.g., the Everett-McNair Store [now Arts
Richmond], formerly badly neglected, has been completely sealed). These renovation and
restoration efforts at buildings, assisted by Cooper’s Hawk predation, have been largely
responsible for the decrease from a moderate to sparse distribution of another urban invader
in Rockingham, the Feral Pigeon.
Recent data, other than imprecise general numbers, are lacking for European Starlings
and House Sparrows in Rockingham, but in many areas of eastern North America since the
bulk of urban and suburban BBCs were conducted in the 1970s and 1980s, urban populations
of House Sparrows have steadily declined (Berigan et al. 2020, Sauer et al. 2017).
This decline includes populations in the Piedmont of North Carolina (Hendrickson and
Ferebee 1994, Westphal 2006). In Sackville, New Brunswick, this formerly abundant species
(Erskine 1980, 1982) is now extirpated (Erskine 2006; A.J. Erskine, retired Canadian
Wildlife Service employee, Sackville, New Brunswick, pers. comm.), and in Gainesville,
Florida, the population has collapsed in suburban areas and sharply declined in urban areas,
where House Sparrows remain most numerous in the central business district (Burnett and
Urban invaders include House Finch populations in eastern North America (Badyaev et al.
2012). This species’ explosive population growth, including in urban areas such as the Piedmont
of North Carolina (Hendrickson and Ferebee 1994, Westphal 2006), was interrupted by
one of the worst epizootics in history—an outbreak of mycoplasmal conjunctivitis. The disease
most likely did not reach Rockingham until between November 1994 and November 1995 (see
Fig. 8 in Dhondt et al. 1998), probably arriving during the latter half of 1995 (Hochachka and
Dhondt 2000). External symptoms of severe conjunctivitis (swollen, crusty, or closed eyes)
were not observed in Rockingham during the breeding season of 1994 (D.B. McNair, pers.
obs.). Thus, this study was conducted when a healthy House Finch breeding population was
near or at its peak in downtown Rockingham in 1994, before its own pandemic began.
2021 No. 44
The resident House Finch occupied a unique position on the Rockingham BBC because
breeding pairs used buildings and vegetation as nest-sites in almost equal proportions (cf.,
Badyaev et al. 2012), yet its number of nest records was only weakly associated with the
area of either buildings or vegetation, especially the latter. This urban invader built its
open-cup nests on, not within buildings, where its distribution was nonetheless restricted to
12 blocks, the same number of blocks as European Starlings and House Sparrows, but the
House Finch was only ~45–50% as numerous as those 2 species. However, House Finches
expanded their distribution within blocks and to additional blocks when they used vegetation
as nest sites. Thus, the House Finch was the most widely distributed avian species that
habitually used buildings on the Rockingham BBC plot, just ~11 years after House Finches
probably first began breeding there ~1982–1983 (LeGrand 1982; LeGrand 1983a,b; Le-
Grand 1984). House Finches may compete with House Sparrows (Cooper et al. 2007, Mc-
Clure et al. 2011), so an assessment of the current status of both urban invaders is urgently
needed to measure changes in their presence or absence and use of buildings and vegetation
as nest-sites within the 25-block area of downtown Rockingham since 1994.
Biotic homogenization in downtown Rockingham worsened between 1994 and the later
period (2012, 2016–2017) from loss of 2 of the 3 most abundant native open-cup species
and a sharp reduction in the number of scarce native species, all of which are categorized as
suburban adapters. The most unexpected result of this study was the loss of breeding Common
Grackles in the 3 later years and the loss, in 2017, of American Robins. Both species
were also absent in 2018 and 2019 (D.B. McNair, unpubl. data). These 2 species formerly
bred at greater heights in vegetation compared to other avian species within the study area
(D.B. McNair, unpubl. data). Almost all of the taller solitary trees, tree rows (including the
former semi-colony site for Common Grackles), most hedges that contain some trees, and
woodlots are still present. Maturation of habitat that has occurred during the 18–23 year gap
between 1994 and the 3 later years would have favored, rather than discouraged, nesting by
these 2 species. Furthermore, extensive lawns, a favorable habitat component for American
Robins, also remain in some blocks. The loss of these 2 omnivores from the central business
district, without replacement by any other abundant native species, has sharply reduced
species richness and abundance on the Rockingham BBC plot.
Breeding Common Grackles are still common in North Carolina (Sauer et al. 2017, Westphal
2009). However, their disappearance from downtown Rockingham is consistent with
a significant long-term decline in the Carolinas since the mid-1960s, which has accelerated
since 2005 (Sauer et al. 2017). This sharp decline includes urban areas of the Piedmont of
North Carolina (Westphal 2009). American Robins first expanded their breeding range to the
Piedmont and Coastal Plain of North Carolina in the 1940s–1950s (Vanderhoff et al. 2016).
American Robins are currently fairly common to common in the Piedmont and southeastern
Coastal Plain (including North Carolina), where they are grouped as an urban breeder (Sauer
et al. 2017). For example, the closest Bird Breeding Survey (BBS) route to Rockingham, at
Hamlet, has averaged 11 American Robins/year since 1995 (range 5–24 birds; Sauer et al.
2017). Robins had a non-significant increase in North Carolina on BBS routes from 1966 to
2015 but decreased by 16% from 2005–2015 (Sauer et al. 2017). No other national or regional
sources have detected a recent decline of breeding American Robins in southeastern North
America. This local study reaffirms the long-term decline of breeding Common Grackles in
North Carolina and also documents the collapse of the American Robin breeding population
in downtown Rockingham, which warrants further investigation.
2021 No. 44
Seven native species that nested as single pairs in 1994 no longer bred in downtown
Rockingham in the 3 later years. The 3 cavity species (Red-bellied Woodpecker, Great
Crested Flycatcher, Brown-headed Nuthatch) visited the study area, but the removal of
scarce snags and loss of nest boxes prevented nesting. Habitat remained substantially unchanged
for the 4 open-cup species (Loggerhead Shrike, Blue Jay [which did visit], Blue
Grosbeak, Orchard Oriole), but they did not breed. Less effort in the 3 later years may have
failed to document a scarce species (but see McNair 2019), but species turnover of scarce
species was substantial despite a rich source of potential immigrants in the immediate vicinity,
including Northern Rough-winged Swallows and Chipping Sparrows (Appendix 1),
which subsequently nested within the Rockingham BBC plot. Absent significant habitat
changes, loss or gain of species that have only occurred as single pairs suggests the central
business district of Rockingham represents sub-optimal habitat. Suburban adapters in the
central business district of Rockingham probably still dominate, but the loss of American
Robins and Common Grackles, which was not caused by vegetative changes, accompanied
by the net loss of scarce native species would reduce suburban adapters’ dominance vis-àvis
exotic urban invaders. Warren et al. (2019) recently documented a loss of bird species,
including suburban adapters, in residential areas of metropoiitan Phoenix, Arizona, that was
also not caused by major vegetative changes. They attributed the loss to an extinction debt,
but the loss of suburban adapters in the old town of Rockingham, where urbanization pressure
is low, is unlikely to be from an extinction debt.
Disregarding standard maintenance practices by municipal and private authorities in the
central business district of Rockingham, the general pattern of benign neglect of vegetative
habitat over the last 20–25 years, except for elimination of isolated snags, has been
favorable for birds in the oldest historical section of this town sitting on top of Cole’s Hill.
Favorable urban typology (Palomino and Carrascal 2006) has allowed Rockingham to live
up to its namesake as a bird sanctuary, despite reduced species richness and bird densities,
which are typical of built-up downtown areas (Cam et al. 2000, Marzluff 2001, Palomino
and Carrascal 2006). Snep et al. (2016) proposed recommendations for habitat enhancement
and other measures that would improve avian biodiversity in urban landscapes across North
America within the context of socio-economic drivers but did not specify any economic
incentives. By North American standards, Rockingham is no longer a wealthy town and
generally lacks an economic impetus to cajole private landowners to improve habitat on
commercial properties beyond minimal standards (City of Rockingham 2020). Thus, the
many recommendations of Snep et al. (2016) are not useful for Rockingham because economic
incentives are lacking.
The town of Rockingham (and Richmond County) also lacks a local cadre of dedicated
field ornithologists to conduct BBCs using the spot-mapping method, unlike Winston-
Salem, North Carolina (Thorington and Brand 2014), a large city. Unlike that study by
Thorington and Brand (2014), other recent studies on urban and suburban birds in southeastern
North America have been conducted by academic biologists using multiple sites and
multi-scaled approaches requiring advanced statistical methods (e.g., at Raleigh and Cary,
North Carolina [Mason et al. 2007]; Gainesville, Florida [Stracey and Robinson 2012]).
Without professional involvement and greater demand, the defenestration of traditional
BBC studies conducted by amateurs is likely to continue unless amateurs couple local
ownership of traditional single-site studies with a long-term approach (e.g., as in Fairfax
County, Virginia; Aldrich and Coffin 1980). Local ownership in Rockingham is lacking,
2021 No. 44
although a local concentration of field ornithologists is present ~50 km away in prosperous
Moore County, North Carolina. As North Carolina’s population continues to rapidly grow
(US Census Bureau 2020), especially in the Piedmont, “local” ownership of single sites
will probably require recruitment of individuals from neighboring communities (McNair
2020). A repeat census of the Rockingham BBC plot, including nest searches, to advance
our understanding of the patterns and processes of biotic homogenization of towns would
be highly desirable. Regardless, future BBCs at most locations that, like Rockingham, are
located in comparatively poor rural areas of southeastern North America will be difficult to
resuscitate without economic incentives.
I thank J.E. Armstrong (past Planning Director, City of Rockingham) and J.R. Massey, Jr. (past
Planning Director and current Assistant City Manager, City of Rockingham) for contributing information;
W. Schmitz (Southeast Regional Climate Center) for directing me to CLIMOD 2; W. Hochachka
(Cornell Lab of Ornithology) for answering 1 query about BBC data; B. Norling for preparing Figure
1 and additional help in performing analyses of geographic information systems data; A.J. Erskine
and F.C. James for their reviews of a penultimate version of the manuscript; and M.W. Strohbach and
2 anonymous individuals for their reviews of the submitted version of the manuscript.
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Appendix 1. List of 50 avian species that visited the Rockingham, North Carolina, breeding bird census plot
from 27 March to 30 July 1994 or nested within 0.4 km outside of its boundary.
Scientific name (Common name) Visitor (V) Nested (N) nearby
Aix sponsa (L.) (Wood Duck) V N
Coccyzus americanus (L.) (Yellow-billed Cuckoo) N
Chordeiles minor (J.R. Forster) (Common Nighthawk) V
Archilochus colubris (L.) (Ruby-throated Hummingbird) N
Butorides virescens (L.) (Green Heron) N
Cathartes aura (L.) (Turkey Vulture) V
Accipiter striatus Vieillot (Sharp-shinned Hawk) V
Ictinia mississippiensis (A. Wilson) (Mississippi Kite) V
Buteo lineatus (Gmelin) (Red-shouldered Hawk) V N
Buteo platypterus (Vieillot) (Broad-winged Hawk) V N
Buteo jamaicensis (Gmelin) (Red-tailed Hawk) V N
Megascops asio (L.) (Eastern Screech-Owl) N
Strix varia Barton (Barred Owl) V N
Megaceryle alcyon (L.) (Belted Kingfisher) V N
Dryobates pubescens (L.) (Downy Woodpecker) V N
Colaptes auratus (L.) (Northern Flicker) V N
Dryocopus pileatus (L.) (Pileated Woodpecker) N
Tyrannus tyrannus (L.) (Eastern Kingbird) N
Contopus virens (L.) (Eastern Wood-Pewee) N
Empidonax virescens (Vieillot) (Acadian Flycatcher) N
Sayornis phoebe (Latham) (Eastern Phoebe) N
Vireo griseus (Boddaert) (White-eyed Vireo) N
Vireo flavifrons Vieillot (Yellow-throated Vireo) N
Vireo olivaceus (L.) (Red-eyed Vireo) N
Corvus ossifragus (A. Wilson) (Fish Crow) V N
Stelgidopteryx serripennis (Audubon) (Northern Rough-winged Swallow) V N
Progne subis (L.) (Purple Martin) V N
2021 No. 44
Apprendix 1. Continued.
Scientific name (Common name) Visitor (V) Nested (N) nearby
Hirundo rustica L. (Barn Swallow) V N
Poecile carolinensis (Audubon) (Carolina Chickadee) N
Baeolophus bicolor (L.) (Tufted Titmouse) V N
Polioptila caerulea (L.) (Blue-gray Gnatcatcher) N
Sialia sialis (L.) (Eastern Bluebird) V N
Hylocichla mustelina (Gmelin) (Wood Thrush) N
Bombycilla cedrorum Vieillot (Cedar Waxwing) V
Spinus tristis (L.) (American Goldfinch) V N
Spizella passerina (Bechstein) (Chipping Sparrow) V N
Zonotrichia albicollis (J.F. Gmelin) (White-throated Sparrow) V
Pipilo erythrophthalmus (L.) (Eastern Towhee) N
Agelaius phoeniceus (L.) (Red-winged Blackbird) N
Molothrus ater (Boddaert) (Brown-headed Cowbird) V N
Geothlypis trichas (L.) (Common Yellowthroat) V N
Setophaga ruticilla (L.) (American Redstart) V N
Setophaga americana (L.) (Northern Parula) N
Setophaga petechia (L.) (Yellow Warbler) V
Setophaga striata (J.R. Forster) (Blackpoll Warbler) V
Setophaga pinus (L.) (Pine Warbler) N
Setophaga coronata (L.) (Yellow-rumped Warbler) V
Setophaga dominica (L.) (Yellow-throated Warbler) V N
Piranga rubra (L.) (Summer Tanager) N
Passerina cyanea (L.) (Indigo Bunting) V N
TOTAL SPECIES 30 41