A systematic review of forest bird occurrence in North American forest fragments and the built environment

Landscape and Urban Planning 185 (2019) 1–23

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Review Article

A systematic review of forest bird occurrence in North American forest fragments and the built environment

T

Jan-Michael J. Archera, , Mark E. Hostetlerb, Glenn Acombc, Robert Blaird ⁎

a

University of Florida, School of Natural Resources and Environment, 110 Newins-Ziegler Hall, PO Box 110430, Gainesville, FL 32611-0430, USA Department of Wildlife Ecology and Conservation, Institute of Food and Agricultural Sciences, University of Florida, Gainesville, FL, USA c Department of Urban and Regional Planning, University of Florida, Gainesville, FL, USA d Department of Fisheries, Wildlife and Conservation, University of Minnesota, Minneapolis, MN, USA b

ARTICLE INFO

ABSTRACT

Keywords: Urban planning Fragmentation Conservation development Landscape design Forest-interior specialist Synthesis

Urban decision-makers typically cannot utilize empirical bird studies to create bird-friendly landscape designs because academic results are often in a format that is not easily transferable to planning. We report on the results of three systematic reviews of peer reviewed literature regarding bird occurrence in forests, forest fragments, and tree canopy in and around North American cities. Note that our reviews focused on occurrence and did not evaluate species fitness or abundance. We synthesized findings from 48 studies to create a list of 219 forest dwelling species; and for each species, we identified whether the species utilized or did not utilize forest fragments during the breeding season; whether migrant species utilized forest fragments as stopover sites during the migration season; and whether the species utilized residential areas with trees during the breeding and migration seasons. We found that some interior-forest specialists, which generally require large tracts of forest to breed successfully, use small urban and rural forest fragments as well as tree canopy within suburban residential areas as stopover sites during migration seasons. This supports previous literature findings that habitat requirements for migrants, including interior-forest specialists, may differ between breeding and migration seasons. The synthesized bird list will allow built environment professionals to identify bird species that can and cannot use treed habitat in fragmented rural and urban landscapes, and we have created a planning tool for conserving avian habitat.

1. Introduction In the wake of rapid urbanization, many researchers have studied how expanding cities impact local and regional bird populations. Generally, ecologists have observed that increasing urbanization negatively impacts native avian species (Aronson et al., 2014). The conversion of forest habitat to urban areas results in fragmentation and loss of native bird habitat; perhaps the biggest direct driver of species’ decline in the context of urbanization (Mckinney, 2006). This is especially true for specialized species that require large areas of forest to successfully breed or lack the functional adaptability to exploit new niches in urban landscapes (Luck & Smallbone, 2010). Urban-rural gradient studies—surveys of population dynamics conducted along transects running from relatively undeveloped city outskirts to heavily built downtown centers—show that native avian species density and richness decrease as development increases (Blair, 1999). Many studies, however, have shown that a wide variety of native

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forest bird species still utilize habitat and breed in heavily developed areas when infrastructure incorporates urban forest fragments or remnants of natural vegetation surrounded by an urban matrix (Donnelly & Marzluff, 2006; Hostetler & Holling, 2000; Hostetler & Knowles-Yanez, 2003). Conserved—or re-planted—areas embedded within cities, such as well-managed riparian buffers or urban parks, safeguard birds against anthropogenic disturbance and have proven to be important urban stopover habitat refugia for many migrating species (Pennington & Blair, 2011; Pennington, Hansel, & Blair, 2008). In addition to conserving forest fragments for use as bird habitat, increasing native tree habitat in residential areas can increase bird diversity in cities (Donnelly & Marzluff, 2006; Hostetler & Holling, 2000). Thus, urban planners can create suitable avian habitat within cities by considering what kinds of natural areas are conserved and by conserving or restoring native vegetation within built areas—neither of which necessarily come at the cost of urban growth and profit (GómezBaggethun & Barton, 2013).

Corresponding author. E-mail addresses: [email protected] (J.-M.J. Archer), [email protected] (M.E. Hostetler), [email protected] (G. Acomb), [email protected] (R. Blair).

https://doi.org/10.1016/j.landurbplan.2019.01.005 Received 21 March 2018; Received in revised form 4 October 2018; Accepted 10 January 2019 0169-2046/ © 2019 Published by Elsevier B.V.

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Despite its numerous potential benefits, evaluating the impacts of different urban designs on avian species and wildlife in general is largely out of reach for decision-makers because this information is accessible only through journal articles and technical reports (Ahern, 2012). Broad conservation strategies, such as conserving large forest tracts, do not fully address the complexities of which species could use remnant habitat when land is subdivided (Hostetler & Holling, 2000; Hostetler & Knowles-Yanez, 2003). Instead, these strategies may overlook more tailored, fine-scale conservation options that can benefit individual species or groups. City planners and developers may be misled to believe that they have few opportunities for conserving bird habitat when land is subdivided. Our goal in conducting this systematic review was to create a reference list for coterminous United States and southern Canadian regions that outlined bird species that were likely to occupy urban or rural forest fragments as well as those bird species that could occupy residential areas with small patches of trees and individual trees (e.g. treed lots, street trees). We sought to answer the following questions: 1.) What bird species occur or do not occur in rural and/or urban forest fragments during the breeding season? 2.) Do certain migrating species use rural and/or urban forest fragments as stopover sites during the fall and spring? 3.) What bird species occur within residential areas with at least partial tree canopy during breeding and migration seasons? Given that research has shown urbanization to have both positive and negative effects on bird populations, these reviews and the resultant bird lists will help built environment professionals to assess alternative development designs and the potential impacts on different bird species. The bird lists could also foster the development of biodiversity planning tools for future planning efforts.

fragment” to be any patch of forest that is less than 50 ha in size and that is surrounded by urban or agricultural landscape. These fragments do not have large built structures under the tree canopy. “Tree canopy in residential area” are urban areas with homes, roads, and scattered trees in the area. We further investigated whether a study indicated if a species preferred early successional forest (i.e., generally composed, mostly, of young hardwood trees, shrubs, grasses, and other species frequently found in regenerating environments) and/or late successional forest (i.e., generally signified by closed-forest canopy and composed mostly of mature conifer and deciduous trees and other slow-growing plant species). For bird habitat, conserving early or late successional forest may affect different bird species. For example, Indigo Bunting (Passerina cyanea) breed primarily along forest edges, which are most often early successional forest habitat (Kohut, Hess, & Moorman, 2009). City decision makers could understand the value of different forest patches if they knew which species would utilize early versus late successional forest. We modeled our systematic review protocol after the methods and framework outlined by the Collaboration for Environmental Evidence (2013). Each review was preceded by compiling a list of Parameter, Indicator, Comparator, and Outcome (PICO) search terms to query the Web of Science© (WoS) online database (Reuters, 2015) (see Supplemental Appendix A1). We defined our geographic scope as studies conducted in Canada and the coterminous United States of America. Our temporal scope was 1970–2015. We also used screening features built into WoS to further exclude certain non-avian studies by Research Domain, Research Area, Document Type, Publication Years, Language, and Countries/Territories (see Supplemental Appendix A2). Authors individually screened resulting articles, rating each as either “0” (not useful) or “1” (potentially useful). Papers were screened at three levels (title, abstract, and full article), using Cohen’s kappa coefficient statistic (Cohen, 1960) to measure inter-rater agreement and ratings were reiterated until we achieved a kappa coefficient greater than or equal to 0.70.

2. Methods 2.1. Systematic review In 2015, we conducted three systematic reviews of peer-reviewed journal articles concerning fragmentation and urbanization impacts on forest birds. We defined “forest birds” as those species that utilize trees, either for foraging or nesting; not solely those species associated with intact forest. Specifically, we selected members of the following taxonomic orders: Passeriformes (commonly known as “perching birds”), Piciformes (woodpeckers), Columbiformes (doves and pigeons), Cuculiformes (cuckoos), and Apodiformes (hummingbirds). We divided our review into three broad categories of habitat use and season: 1) bird studies of rural forests and urban forest fragments used as breeding habitat during the summer season (hereafter referred to as the Breeding Review), 2) bird studies of urban and rural forest fragments used as stopover habitat during fall and spring migration seasons (hereafter Stopover Review), and 3) bird studies within city, residential areas that contain trees, during breeding and migration seasons (hereafter Built Environment Review). We used these habitat categories because the same avian species have been recorded to use different habitats throughout the year (e.g., Hostetler & Holling, 2000). For example, interior-forest specialists (e.g. Hooded Warbler Wilsonia citrina) exhibit area-sensitivity during the summer breeding season, generally requiring large contiguous areas of mature or late successional forest (Devictor, Julliard, & Jiguet, 2008). During fall and spring migration seasons, though, some of these interior-forest specialists show little area sensitivity and often utilize small forest fragments as stopover sites along flyways (Seewagen, Slayton, & Guglielmo, 2010). Also, city planners make different decisions when trying to conserve urban forest fragments versus trees and tree canopy in residential areas. For this review, we consider a “forest

2.2. Avian species lists 2.2.1. Breeding review From studies conducted in urban and rural forest areas during the breeding season, we assigned bird species to one of two habitat-usage groups: interior-forest specialists (those species requiring core interior forest habitat), and non-interior-forest specialists (those species utilizing regenerating early successional habitat at the forest’s edge or habitat throughout the forest). Our main goal was to determine which species were consistently considered interior-forest specialists and would not use urban or rural forest fragments for breeding versus those that would. We selected studies that: a.) conducted bird surveys across a range of forest fragment sizes and correlated species density/presence with fragment size (e.g., Ambuel & Temple, 1983), b.) conducted bird surveys along a transect from a forest’s edge to its interior and listed which species occurred relative to position along this transect (e.g., Blake & Karr, 1984), and c.) determined area-sensitivity of a species and determined correlations between species density and increasingly fragmented landscapes (e.g., Boulinier et al., 2001). Interior-forest specialists were defined here as those occurring either exclusively in large forest patches (> 50 ha) or explicitly occurring within the core forest (as opposed to edge forest). Species that were not interior-forest specialists met one of two conditions: 1) they occurred throughout patches (sometimes referred to as “interior-edge” in the literature) and whose density showed no significant correlations with forest patch size or degree of landscape fragmentation, or 2) they

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Fig. 1. Breeding Review Decision Hierarchy. For this review, species were considered not to be interiorforest specialists only if assigned low confidence of being interior-forest specialists or no studies stated that it was an interior-forest specialist. High- and medium-confidence interior-forest specialists were interior-forest specialists in this review.

occurred on the edge of patches (sometimes referred to as “edge” species in the literature) and whose density decreased with increased forest patch sizes. To deal with conflicts between studies (e.g. where one paper found a particular species to be an interior-forest specialist but another did not), we ranked overall confidence in a particular classification based on the following (Fig. 1): For species recorded in three or more papers, if > 67% of papers indicated the species was an interior-forest specialist, we reported high confidence that it was an interior-forest specialist. If 34–67% of the studies indicated the species was an interior-forest specialist, we reported medium confidence. If ≤33% of articles indicated a species was an interior-forest specialist, we reported low confidence. Only high/medium confidence levels indicated that a species was an interior-forest specialist. If a species had a low confidence level, or it was reported in fewer than three papers, we simply reported how many papers indicated that a species was an interior-forest specialist, but we did not designate it as an interior-forest specialist. If more than three studies were conducted and none of them indicated that the species was an interior-forest specialist, we reported that it was not an interior forest specialist. Next, we distinguished between birds that prefer to breed in late- or early successional forest in fragmented landscapes. We referred to life history data from the Cornell University Lab of Ornithology’s All About Birds© website (www.allaboutbirds.com) and The Birder’s Handbook: A Field Guide to the Natural History of North American Birds (Ehrlich, Dobkin, & Wheye, 1988) to assign whether birds breed in early successional forest, late successional forest, or both. The rationale for distinguishing between early and late successional breeding birds is

that, when creating an urban design, conserving late vs early successional forest will impact different species. For example, woodpeckers primarily breed in large trees found in late successional forests (Ehrlich et al., 1988); if a woodpecker species was tolerant of fragmentation, it would primarily be found in late successional forest fragments. 2.2.2. Stopover review We concentrated on studies that explored migrants and their use of urban and rural forest fragments as stopover sites. We compiled a list of reported migratory species surveyed in small fragments (less than 50 ha) during the spring and fall migration seasons. After referring to All About Birds© and Ehrlich et al. (1988), we confirmed each species’ migrant status and whether they were long-distance, medium-distance, short-distance, or irruptive. We defined “long distance” as intercontinental migration (e.g., from Canada and the United States to Central and South America), “medium-distance” as trans-regional migration (e.g. from Southern Canada to Mid-Atlantic United States), and “short-distance” as within-region migration (e.g. from one state to a neighboring state). Irruptive migrations are those occurring unpredictably, usually because of sudden resource limitation. Some species utilize a combination of migration strategies. For example, the Yellow-rumped Warbler Setophaga coronata, have both medium-distance migrants (migrating to south Florida from Canada) and longdistance migrants (migrating to South America). We reported this as well. As mentioned in the previous section, the occurrence of most breeding interior-forest specialist populations has a strongly positive relationship with increasing sizes of forest fragments. To identify the

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extent to which migrant birds might exhibit this effect during stopover, we made note of studies that compared species occurrence during the fall and spring migration seasons across relatively small forest fragments that varied in size. This information could indicate whether migrants, in general, tend to select relatively larger forest fragments in a landscape.

2.2.5. Overall comparisons If a species was recorded in studies across the three reviews or in two or just one review category, we evaluated whether certain species would or would not occur in urban/rural forest fragments or in residential areas. For ease of interpretation, we created four “Overall” columns in our final bird list: “Breeds in Forest Fragment”; “Stopover in Forest Fragment”; “Breeds in Residential Area”; and “Stopover in Residential Area.” For a given species, a checkmark (✓) indicates its likely occurrence in a habitat category, an (X) indicates that it does not occur in this habitat category, a (?) indicates that a species may occur in this habitat category but results were not convincing enough to assign either a (✓) or (X), and cells left blank indicate that no information was available. Species identified solely in Breeding Review studies and were given a High/Med confidence level as an interior-forest specialist, we assigned an (X) in the “Breeds in Forest Fragment” and “Breeds in Residential Area” categories. This signifies that the species does not breed in forest fragments in rural and urban areas. Species with Low confidence or observed by a total of two or fewer studies in the Breeding Review, we assigned a (?) to indicate that this species may or may not breed in forest fragments in urban and rural areas. Species that had three or more studies that indicated that it was not an interiorforest specialist was assigned a (✓), and it most likely breeds in forest fragments. Migrant species observed by three or more studies but solely in the Stopover Review were assigned (✓) under the “Stopover in Forest Fragment” category. If a migrant had fewer than three studies in the Stopover Review, then we assigned (?) under the “Stopover in Forest Fragment” category. For migrants found in both the Stopover Review and Built Environment Review, if Stopover Review studies combined with two or fewer Built Environment Review studies (stopover, breeding, and breeding-stopover studies), which brought the total to three or greater, then we assigned a (✓) for Stopover in Forest Fragment and a (?) for Stopover in Residential Area categories. The rationale here is that migrants seen in two or fewer Built Environment studies receive a (?) for Stopover in Residential Area because that is not enough studies to confidently assign them as using residential areas as stopover sites. For species found in the Breeding Review, Built Environment Review, and Synanthropy Analysis, we compared results to determine what to assign in each habitat category. If a species was a High/Medium confidence interior-forest specialist but was observed in three or more studies in the Built Environments Review during the breeding season and was shown to have at least one study indicating it was an urban dweller, we assigned a (✓) for this species as Breeding in Forest Fragment and Breeding in Residential Area. If a migrant, we also assigned a (✓) for “Stopover in Forest Fragments” and “Stopover in Residential Area” because if it breeds in residential areas we assumed it would use urban forest fragments and trees in residential areas as stopover sites. We understand that there is no empirical evidence for these migrants to use stopover areas in residential and in forest fragments; however, it stands to reason that a bird that is breeding in a highly fragmented residential area, it would not be averse to using residential or forest fragment areas as stopover sites. Exceptions could occur and thus we marked the (✓) with an “*” to indicate that this check is not based on empirical evidence for the given habitat category. For species found in Breeding Review, Built Environment Review, but not in the Synanthropy Analysis, a High/Medium/Low or unassigned confidence level for interior-forest specialist that was a yearround (YR) resident and/or a short-distance (SD) migrant, and it occurred in three or more Built Environment breeding studies, we assigned a (✓) as “Breeding in Forest Fragment” and “Breeding in Residential Area.” We were confident that the residential studies

2.2.3. Built environment review From studies conducted within urban and suburban residential areas, we created a list of reported species sighted during both breeding and migration seasons, investigating whether seasonality played a role in species occurrence within the built environment. Particularly, we hoped to highlight that while a species may not breed within a residential area, it may still use tree canopy and associated vegetation as a stopover site. For studies that did not adequately report when a survey occurred (e.g., study periods that ran from spring into summer or from summer into fall) we recorded the survey period as “BreedingMigration.” Note that long-distance migrants that were recorded in these studies were most likely using the residential area as a stopover site. Also, studies that were recorded as breeding surveys were typically started in May/June, but May/June could also have migrating birds stopping over in residential areas. Thus, all long-distance and mediumdistance migrating birds occurring in residential areas during the breeding season may or may not be breeding in residential areas; we chose to be conservative and not assign these species as breeding in residential areas. We focused on reviewing research that studied birds primarily utilizing trees and shrubs. The amount of vegetation coverage represents a key component of urban landscape design at the residential scale (e.g., Hostetler & Holling, 2000). We noted studies that investigated relationships between species occurrence and tree canopy cover as well as relationships between occurrence and native versus exotic trees. 2.2.4. Synanthropy analysis From our three systematic reviews, we identified four studies that investigated synanthropy—a measure of tolerance to urbanization. We defined synanthropic species as those that occurred in forest fragments or residential areas and whose density or presence was positively correlated with increasing amounts of surrounding urban land use. We update this terminology to use the term “urban dweller” that is equivalent to a synanthropic species (Fischer, Schneider, Ahlers, & Miller, 2015). Species that are not urban dwellers were defined as those occurring in forest areas or residential areas that were negatively correlated with increasing amounts of urban land use. The four studies were conducted along urbanization gradients and were: Donnelly and Marzluff (2006); Galitsky and Lawler (2015); Oneal and Rotenberry (2009); and Parrish and Hepinstall-Cymerman (2012). Donnelly and Marzluff (2006) conducted their study at the 1 km landscape scale (including bird surveys in both residential areas and forest fragments surrounded by residential areas) while the other three studies surveyed birds exclusively in forest fragments within cities. Providing a list of urban dweller and non-urban dweller species allows urban decision-makers to assess how local bird populations inhabiting forest fragments may change after surrounding development is increased. Where conflicts occurred for species that had at least three studies, we gave confidence of a species being an urban dweller in the same way that we assigned confidence in the Breeding Review (i.e., ≤33%, 34%–67%, and > 67% papers reporting a species was an urban dweller related to low, medium, and high confidence, respectively). For species recorded in fewer than three studies, we simply noted how many studies showed that a species was an urban dweller.

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reflected breeding individuals because the birds were not migratory. If a High/Medium or unassigned confidence level for interior-forest specialist was a long-distance and/or medium-distance migrant species (even if a portion of the population is considered year-round) and occurred in three or more Built Environment breeding surveys, we were conservative and assumed these individuals were passing through cities. We assigned a (✓) for these species under “Stopover in Residential Area” and “Stopover in Forest Fragments.” However, if a migrant species was determined to be a Low-confidence interior-forest specialist and occurred in three or more Built Environment Review breeding studies, we assigned a (✓) for the “Breeds in Residential Area” and “Breeds in Forest Fragment” categories because the combination indicates that they may also breed in residential areas. In turn, because they breed in residential areas, then they would also tolerate fragmented forests. If a bird breeds in a residential area, we assumed it would breed in a forest fragment. Again, we marked the (✓) with an “*” to indicate that this check is not based on empirical evidence for the given habitat category. For species that occurred solely in the Built Environment Review and had three or more total studies (combined stopover, breeding, and breeding-stopover studies), we interpreted the likelihood of this species breeding in residential areas in the following way: If they were observed within the built environment during the breeding season and were shown to be an urban dweller in at least one synanthropic study, then we assigned a (✓) for “Breeding in Forest Fragment” and “Breeding in Residential Area.” If a migrant, we also assigned a (✓) for “Stopover in Forest Fragments” and “Stopover in Residential Area” because if they breed in residential areas they would use forest fragments and residential areas as stopover sites. If species were only observed in the breeding season surveys (not in stopover and breedingstopover studies), we assigned a (✓) only for YR residents and SD migrants under “Breeds in Forest Fragment” and “Breeds in Residential Area” categories. All long-distance and medium-distance migrants were considered individuals that were passing through, most likely using the areas as a stopover site. Here, we assigned a (✓) for these species for “Stopover in Forest Fragment” and “Stopover in Residential Area” categories. Additionally, for long-distance and medium-distance migrants that had three or more combined stopover, breeding, and breedingstopover studies under the Built Environment Review, we assigned a (✓) for these species for “Stopover in Forest Fragment” and “Stopover in Residential Area” categories. If a migrant used stopover sites in residential areas, we assumed it would also use stopover in forest fragments because of its tolerance of highly fragmented residential areas. Again, for these situations, we marked the (✓) with an “*” to indicate that this check is not based on empirical evidence for the given habitat category. For species that occurred solely in the Built Environment Review and had fewer than three total studies, we interpreted the likelihood of this species breeding in residential areas in the following way. For YR resident species and/or SD migrants, we assigned a (?) in the corresponding “Breeds in Forest Fragment” and “Breeds in Residential Area” categories. All long-distance and medium-distance migrants were assumed to be passing through and may be using residential areas as stopover sites. We assigned a (?) for these species for “Stopover in Forest Fragment” and “Stopover in Residential Area” categories.

reduced the number of usable studies from 2612 to 18. In the Built Environment Review, screening reduced the number of usable studies from 1893 to 18 (Table 1). From these studies, we compiled lists of forest birds (219 species total) where we marked interior-forest specialist status, migrant status, early or late successional forest preference, and tolerance to urbanization (Appendix A). Of these 219 species, 156 were observed in at least two of our three reviews, allowing us to compare results from breeding and migration seasons and from studies conducted within the built matrix. 3.2. Avian species lists 3.2.1. Breeding review The systematic review of peer-reviewed literature on forest fragments as avian breeding habitat generated a total of 81 forest birds. We recorded species as having “high” (14), “medium” (6), and “low” (5) confidence in being interior-forest specialists, respectively. Eleven species were assigned “no” confidence of being interior-forest specialists (Appendix A). There were 45 species (of the 81-total species) that were observed by fewer than three studies, a low sample size for our purposes. These species were not assigned any confidence score. All 20 species with high or medium confidence of being an interiorforest specialist, per All About Birds ©, inhabit late-successional forest habitat, which included the Yellow-throated Vireo (Vireo flavifrons) that inhabits the edges of late-successional forest habitat. Long-distance migration was the most represented migration type among these interior-forest specialists with 16 species (e.g., Ovenbird Seirus aurocapilla) exhibiting it. We listed low confidence of being an interior-forest specialist for these five species: American Crow (Corvus brachyrhynchos), Baltimore Oriole (Icterus galbula), Great Crested Flycatcher (Myiarchus crinitus), Tufted Titmouse (Baeolophus bicolor) and Yellow-billed Cuckoo (Coccyzus americanus). Both the Yellow-billed Cuckoo and Great Crested Flycatcher exhibit long-distance migrations and prefer open woodland habitat while the Tufted Titmouse is a year-round resident that breeds in forests. Finally, 11 species were not observed to be interior-forest specialists by any study. This included many common birds such as the American Robin (Turdus migratorius), Northern Cardinal (Cardinalis cardinalis), European Starling (Sturnus vulgaris), Blue Jay (Cyanocitta cristata), Black-capped Chickadee (Parus atricapillus), and Downy Woodpecker (Picoides pubescens). Notably, while most of the species in this last group are year-round residents or short-distance migrants, the Eastern WoodPewee (Contopus virens) and Indigo Bunting (Passerina cyanea) are both long distance migrants. 3.2.2. Stopover review The systematic review of peer-reviewed literature on forest fragments as avian stopover habitat generated a list of 117 migrant species (SD, MD, or LD) that used small forest fragments as stopover sites (Appendix A). Studies measured forest bird occurrence in fragments that ranged from < 1 to 20 ha. Most species observed in this review were long-distance migrants (n = 77). There were only four species known to exhibit irregular migrations in this list. Of the 18 studies used in our review of stopover habitat, four offered comparisons of species presence between larger and smaller forest fragments: Cox (1988); Somershoe and Chandler (2004); Keller and Yahner (2007); and Suomala, Morris, Babbitt, and Lee (2010). In Cox (1988), 10 of the 54 species were found exclusively in fragments larger than 20 ha in area; the rest were found in fragments of area less than 5 ha. In Somershoe and Chandler (2004) regression analyses showed significant positive relationships between increasing forest fragment sizes—ranging from 0.32 to 3.08 ha—and densities in American

3. Results 3.1. Systematic review Initially, our WoS database query returned 7511 studies across the three systematic reviews. In the Breeding review, screening reduced 3006 publications to 12 studies. In the Stopover Review, screening

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Table 1 Journal Articles Used for Synthesis. From three systematic reviews, forty-eight studies were identified and used in the generation of bird lists. Four studies, marked by double asterisks, were used to determine a synanthropic rating for some species of birds. Source

Journal

Study Location

Review

Ambuel and Temple (1983) Austen, Francis, Burke, and Bradstreet (2001) Blake and Karr (1984) Blake and Karr (1987) Boulinier et al. (2001) Chan and Ranganathan (2005) Galitsky and Lawler (2015)** Galli, Leck, and Forman (1976) Howell et al. (2000) Lapin, Etterson, and Niemi (2013) Richmond, Nol, and Burke (2012) Robbins et al. (1989) Cox (1988) Liu and Swanson (2014)

Ecology The Condor Biological Conservation Ecology Ecology Oikos Landscape Ecology The Auk Landscape Ecology The Condor Canadian Journal of Zoology Wildlife Monographs Florida Field Naturalist Physiological and Biochemical Zoology Landscape Ecology The Condor The Wilson Bulletin The Auk The Auk The Auk Southeastern Naturalist Northeastern Naturalist Wilson Journal of Ornithology Acta Oecologica – Int’l Journal of Ecology The Wilson Bulletin Wilson Journal of Ornithology American Midland Naturalist The Wilson Bulletin The Wilson Bulletin The Auk Ecological Applications Conservation Biology Urban Ecosystems Ecological Applications

Pennsylvania, USA Ontario, Ca Illinois, USA Illinois, USA BBS* Ontario, Ca Oregon, USA New Jersey, USA Missouri, USA Minnesota, USA Maryland/ Pennsylvania/ West Virginia/ Virginia, USA Ontario, Ca Florida, USA South Dakota, USA

Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Stopover Stopover

Ohio, USA Ohio, USA Pennsylvania, USA Pennsylvania, USA Pennsylvania, USA Pennsylvania, USA Tennessee, USA New York, USA New York, USA New York, USA

Stopover Stopover Stopover Stopover Stopover Stopover Stopover Stopover Stopover Stopover

South Carolina, USA Maine/New Hampshire, USA South Dakota, USA Minnesota, USA Minnesota, USA Alabama, USA Chicago, Illinois, USA Southeastern Pennsylvania, USA Seattle, Washington, USA Tucson, Arizona, USA

Stopover Stopover Stopover Stopover Stopover Stopover Built Environment Built Environment Built Environment Built Environment

Landscape and Urban Planning Landscape and Urban Planning

Phoenix, Arizona, USA Amherst/Springfield, MA; Austin, TX; Blacksburg, VA; Chicago, IL; Seattle, WA; Vancouver, B.C. Phoenix, Arizona, USA Gainesville, Florida, USA Raleigh/Cary, North Carolina, USA Phoenix, Arizona, USA Chicago, Illinois, USA Sonoma County, California, USA Tucson, Arizona, USA Tucson, Arizona, USA Baltimore, Maryland, USA Orange County, California, USA Athens, Georgia, USA Lake Tahoe Basin, California/Nevada, USA

Built Environment Built Environment

Matthews and Rodewald (2010a) Matthews and Rodewald (2010b) Rodewald and Brittingham (2002) Rodewald and Brittingham (2004) Rodewald and Brittingham (2007) Rotenberry and Chandler (1999) Rush, Soehren, and Miller (2014) Seewagen (2008) Seewagen and Slayton (2008) Seewagen et al. (2010) Somershoe and Chandler (2004) Suomala et al. (2010) Swanson, Carlisle, and Liknes (2003) Weisbrod, Burnett, Turner, and Warner (1993) Winker, Warner, and Weisbrod (1992) Woodrey and Moore (1997) Belaire et al. (2014) Burghardt et al. (2009) Donnelly and Marzluff (2006)** Germaine, Rosenstock, Schweinsburg, and Richardson (1998) Green and Baker (2003) Hostetler and Holling (2000) Hostetler and Knowles-Yanez (2003) Hostetler, Duncan, and Paul (2005) Kohut et al. (2009) Lerman and Warren (2011) Loss et al. (2009) Luther et al. (2008) McCaffrey and Mannan (2012) Mills et al. (1989) Nilon, Warren, and Wolf (2009) Oneal and Rotenberry (2009)** Parrish and Hepinstall-Cymerman (2012)** Schlesinger et al. (2008) *

Landscape and Urban Planning Southeastern Naturalist Urban Ecosystems Ecological Applications Biological Conservation Biodiversity and Conservation Landscape and Urban Planning The Condor Urban Habitats Landscape and Urban Planning Urban Ecosystems Ecology

Built Built Built Built Built Built Built Built Built Built Built Built

Environment Environment Environment Environment Environment Environment Environment Environment Environment Environment Environment Environment

The Breeding Bird Survey (BBS) includes study sites across the United States and Southern Canada. These studies were used for our Synanthropy Analysis.

**

Redstart, Blue-gray Gnatcatcher, Eastern Wood-pewee, Northern Waterthrush, and Palm Warbler presence. Suomala et al. (2010)—using only two study islands—determined that eleven migrant species on Appledore Island, Maine (33.6 ha in area) did not occur on Star Island (13.4 ha in area). In Keller and Yahner (2007), several migrants were more likely to occur in larger forest patches than in smaller patches during the spring-migratory season but not the fall-migratory season. Overall, it seems that some migrants are more likely to use relatively larger patches of forest as stopover sites.

3.2.3. Built environment review From the 18 studies of forest birds within the built environment, we listed 201 avian species observed within urban residential areas during the breeding and migration seasons (Appendix A). Studies were conducted primarily in fragmented areas dominated by single family homes (1–7 homes/hectare), containing some multi-family housing and commercial properties. Of the 201 species observed, 143 were in residential areas during the breeding season, 90 species were observed in residential areas during the spring or fall migration seasons, and 99

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species were observed during the “Breeding-Migration” survey period. Three studies in our review did not clearly state whether observations occurred during the breeding or migration season and we listed species from these papers into a “Breeding-Migration” column (Appendix A). There were 56 long-distance migrant species observed during the migration seasons, indicating that these species were utilizing residential areas containing tree canopy as stopover habitat. Overall, the most reported species occurring in the built environment were common species such as the House Finch (Carpodacus mexicanus) (n = 13 studies), Brown-headed Cowbird (Molothrus ater) (n = 12), Northern Cardinal (Cardinalis cardinalis) (n = 12), Mourning Dove (Zenaida macroura) (n = 11), and Northern Flicker (Colaptes auratus) (n = 11). From our review, articles that studied the effect of canopy cover on local bird communities within the built matrix found that an increase in canopy cover resulted in an increase in forest bird diversity: Belaire, Whelan, and Minor (2014); Donnelly and Marzluff (2006); Green and Baker (2003); Kohut et al. (2009); and Luther et al. (2008). The following five studies from our review investigated the effects of native vegetation and found that more native birds were present when more native vegetation was in the area: Burghardt, Tallamy, and Shriver (2009); Lerman and Warren (2011); McCaffrey and Mannan (2012); Mills, Dunning, and Bates (1989); and Schlesinger, Manley, and Holyoak (2008).

two species also use forest fragments and residential areas as stopover sites because they occurred in studies conducted during migration season in forest fragments and in residential areas. Our overall comparisons revealed habitat use for many species during breeding and migration seasons (Appendix A). We broke this down according to whether a species had year-round (YR) and/or shortdistance (SD)/irruption (IR) migration status. Eliminating the one interior-forest specialist (i.e., Pileated Woodpecker r Hylatomus pileatus), we identified 86 species of year-round residents (YR) and/or short distance (SD)/irruption (IR) migrants that had no long-distance (LD) or medium-distance (MD) designation. Of these, 45 (52%) species could breed in both forest fragments and residential areas (i.e., “✓”) and one species had a “✓” in forest fragment category only. Of the rest, 40 (47%) had at least a “?” in either the forest fragment or residential review column or both. Thus, it seems YR/SD designation meant a species was more likely to breed in fragments. We also identified 107 species of MD and/or LD migrant species that did not have a YR or SD designation. Fifteen species (14%) could breed in both forest fragments and residential areas and 9 species (8%) could breed in forest fragments only. Of the rest, 56 of 107 (52%) had a “blank” for both forest fragment or residential column and 27 of 107 (25%) had at least one “?” For migration studies, 22 of 38 species of SD and/or IR only migrants (58%) could use both fragmented forest and residential areas for stopover habitat, 2 species (5%) had a “✓” for forest fragment category only, and 14 (37%) had a “?” for either category. For MD and/or LD only migrants, 46 of 107 (43%) could use both forest fragments and residential areas as stopover sites. Of the rest, 28 of 107 (26%) had a “✓” for forest fragment and a “?” for residential studies and 33 of 107 (31%) studies had a “?” for forest fragment and/or residential. Thus, it seems that a majority of short- and long-distance migrants can use forest fragments and even residential areas as stopover sites. Interestingly, of the 15 interior-forest specialists designated in the overall comparison, which included 14 long-distance migrants, 10 migrants did use urban and rural small fragments and/or residential areas as stopover sites.

3.2.4. Synanthropy analysis From the three systematic reviews, we found four synanthropic studies and, from these, compiled a list of 114 bird species. Of the 20 species found in three studies or more, nine were assigned high confidence of being an urban dweller species, six were assigned medium confidence, one was assigned low confidence, and four were assigned no confidence (Appendix A). Forty-nine species were not listed as urban dwellers by any of the four studies. Notably, Hutton’s Vireo (Vireo huttoni), Pacific-slope Flycatcher (Empidonax difficilis), Spotted Towhee (Pipilo maculatus), and Wilson’s Warbler (Wilsonia pusilla), were each observed by three of the four studies in our analysis but not found to be urban dwellers by any of them. As such, we have highest confidence that these four species are not urban dwellers.

4. Discussion

3.2.5. Overall comparisons As mentioned earlier, of the 219 species, 156 were observed in at least two of our three reviews. There were 60 species in the Breeding Review that were also observed in the Built Environment Review and some of these were in the Synanthropic Analysis as well. After comparing these observations, we assigned each species a (X), (✓), or a (?) under each of four occurrence categories: “Breeds in Forest Fragment”; “Stopover in Forest Fragment”; “Breeds in Residential Area”; and “Stopover in Residential Area.” (Appendix A). For example, Red-bellied Woodpecker (a year-round resident) and Red-eyed Vireo (long distance migrant) had medium confidence levels for interior-forest specialist, but they occurred in three or more studies under the Built Environment Review and in at least one study that indicated they were an urban dweller species. The results of combining these studies helped us to designate that the Red-bellied woodpecker would probably breed in forest fragments and residential areas and the Red-eyed Vireo would probably breed in forest fragments and residential areas and use them as stopover sites. Overall, the 20 designated interior-forest specialists (from Breeding Review) dropped to 15 because of the overall review. As an example, for two of the five species that were dropped, two mediumand long-distance migrant species, Baltimore Oriole (Icterus galbula) and Great Crested Flycatcher (Myiarchus crinitus), were determined to possibly breed in fragments because these were assigned low confidence as an interior-forest specialist but had more than three observations in the built environment during the breeding season. These

4.1. Systematic review Our reviews yielded data from 48 peer-reviewed studies that allowed us to compile a list of 219 species of forest birds, all of which utilized small forest fragments and trees within residential areas in some way either during the breeding or migration season or both. Our screening process was intentionally strict to find papers that analyzed individual species. The review was robust as we were able to compare across bird studies in urban/rural fragments and studies in residential areas. There are, however, some limitations to this review. First, we only collected English-language publications from the Web of Science online publication database, giving our data an inherently AngloAmerican bias. Secondly, there was very little representation of studies conducted in the western portion of North America—only 12 publications of the total 48 used in the reviews. Further, certain species not observed in the Built Environment or Stopover Reviews may not have breeding ranges or migration flyways that overlap with the locations of the studies. We also acknowledge that members of the same species may select different breeding and foraging habitats across the geographic scope of our study (e.g. American Redstart Setophaga ruticilla foraging along shorelines utilize more coniferous trees than their inland counterparts that utilize deciduous trees) (Smith, Hamas, Ewert, & Dallman, 2004). However, we believe the sampling scope of our study is enough to provide a general indication of whether a species uses forest fragments and/or residential areas during the breeding and

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migration seasons. Despite its limitations, this systematic review will help planners determine the impacts of development designs and city designs on forest birds. Below we discuss our findings in the context of city and county planning design decisions.

forest fragment contains disturbed and regenerating early-successional forest along the edge, unless it was recently clearcut. The outermost edges of forest patches generally sustain more disturbance (from both anthropogenic and natural sources) than the forest’s interior core. Thus, these edges are composed mostly of early-successional, shrubland habitat. Several so-called “edge specialists” such as the Indigo Bunting, Song Sparrow, and Yellow-breasted Chat are more accurately described as shrubland specialists because they will make use of shrubland habitat, even if it occurs within a greater forested matrix, e.g., where there has been a clearcut or blowdown or other disturbance that gives way to regeneration (Forsman, Reunanen, Jokimäki, & Mönkkönen, 2013).

4.2. Avian species lists 4.2.1. Forest fragments as breeding habitat for birds We identified 15 species that we assigned as overall interior-forest specialists, including species like the Cerulean Warbler (Dendroica cerulea), a Neotropical migrant listed as vulnerable (i.e. likely to become endangered) by the International Union for Conservation of Nature (IUCN) (Birdlife International. (2015), 2015). Thus, during breeding season, these 15 species will not select urban or fragmented landscapes containing small forest patches. First described in the 1980s, interiorforest specialists generally show an aversion to habitat fragmentation (Cox, 1988; Robbins, Dawson, & Dowell, 1989). For interior-forest specialists, fragmented landscapes caused by urbanization and agriculture generally do not provide high quality breeding habitat due to increased nest predation and to parasitism by cowbirds (Ibáñez-Álamo et al., 2015), competition with invasive species (Shochat et al., 2010), and anthropogenic pollution (e.g. noise, light, chemical, etc.) (Laurance, 2008). Even when these remaining fragments represent good quality habitat, the break in connectivity poses problems for some birds because traversing roads, bridges, open fields, and other humanmade structures involves high risk of predation and collision (Loss, Ruiz, & Brawn, 2009). At regional scales, by preserving large forested areas outside of cities and agricultural areas, planners can protect breeding habitat of species threatened by fragmentation. Additionally, these preserved forests sustain populations of habitat generalists, keeping common species, like the Northern Cardinal, common. Using our overall comparisons between the Breeding Review, the Built Environment Review, and Synanthropic analysis revealed discrepancies in the literature. Five species originally assigned High and Medium confidence of being interior-forest specialists were also reported to reside within residential areas and had a high confidence of being urban dwellers during the breeding season. Ultimately, these five species were not considered to be interior-forest specialists. Additionally, we originally assigned the American Crow with low confidence of being an interior-forest specialist in our Breeding Review because it was observed to be so by one of three studies (Boulinier et al., 2001). The results of our Synanthropic analysis, though, confirmed that the American Crow is indeed an urban dweller. As another example, the Red-eyed Vireo was assigned medium confidence as an interior-forest specialist, but it was reported in three built environment studies, one of which indicated that it was an urban dweller. For urban design to attract certain birds that could breed in fragmented landscapes, one must also note the type of forest fragment retained – is it late or early successional forest? For instance, the Whitebreasted Nuthatch nests almost exclusively in cavities of large, mature trees (Wiebe, 2011). A late-successional forest fragment provides considerably more nesting opportunities than an early-successional forest with much smaller trees. White-breasted Nuthatch was also found to breed in residential areas, indicating that mature coniferous and deciduous trees, such as pines and oaks in neighborhood yards or along streets, represent prime nesting sites for this species. This goes for other cavity nesters, such as woodpeckers, titmice, and flycatchers, as well. Thus, developers that conserve late successional forest fragments with large trees and individual large trees in residential areas promote breeding habitat for these species. In urban planning context, preserving late-successional forest fragments also means preserving early-successional habitat, increasing the densities of both generalist and edge specialist species. Each mature

4.2.2. Fragments as stopover habitat for forest birds Our review of forest fragment habitat used during migration seasons revealed that 117 species of migrant birds utilize forest fragments as stopover sites. Interestingly, 10 of the 14 species identified (overall) as interior-forest specialist migrants utilized both urban and rural forest fragments and residential areas during fall and spring migration seasons. This agrees with previous studies’ findings that while forest fragments in urban and rural areas are not conducive for breeding, urban/rural forest fragments may be important for migrating birds as stopover habitat (Woodrey & Moore, 1997). Even residential areas with trees can provide stopover habitat for many interior-forest specialist migrants as discovered in our review. Additionally, most species observed in this review were long-distance migrants, indicating that these forest fragments play a role supporting bird populations throughout North America. Such populations are protected by trans-national legislation such as the Migratory Bird Treaty Act of 1918 because they cross international boundaries. Though it was infeasible to determine a minimum patch size for all interior-forest specialists, four of the studies in our Stopover Review indicated that relatively larger forest fragments contained more migrating individuals or more migrant species than smaller fragments (Cox, 1988; Keller & Yahner, 2007; Somershoe & Chandler, 2004; and Suomala et al., 2010). It may be that larger forest fragments are more attractive to migrants because they present more foraging opportunities. Matthews and Rodewald (2010a) measured within-patch movements of Swainson’s Thrushes (Catharus ustulatus) at stopover sites ranging in size from 0.7 to 38.4 ha in New York City, New York. These researchers found that individuals released into the two smallest study sites (0.7 ha and 4.5 ha) relocated almost immediately while individuals released into larger study sites (the next tier being 11 ha) did not relocate at all. From our review, it was impossible to get a minimum patch size as the studies used various patch ranges; although all of them indicated relatively larger patch sizes had more species or increased abundances, smaller patches may have limited resources and only attractive to a few migrants. While 1.0–5.0 ha patches can serve as stopover sites, if options exist, perhaps planners should look for ways to conserve relatively larger forest fragments (> 5.0 ha) for migrant species. Still, small forest fragments can represent very suitable stopover habitat for migrants. Matthews and Rodewald (2010b) also observed that Swainson’s thrushes showed no trends in stopover duration (the amount of time spent at a stopover site during migration) between release sites along an urban-gradient in Columbus, Ohio—despite the sites varying in size (from 11.9 ha to 38.4 ha) and level of urbanization surrounding patches. Seewagen et al. (2010) found that Ovenbird (Seiurus aurocapillus), a high-confidence interior-forest specialist species with several observations in our review, utilized fragments as small as 1 ha in Prospect Park, New York City, USA. It is worth noting that this park, though deeply embedded within the urban built environment matrix, is directly along the Atlantic Flyway and, thus, may represent an outlier in migrant visitation. Liu and Swanson (2014) studied

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refueling in several migrant birds that utilized either anthropogenic farmstead woodlots or naturally wooded corridors as stopover habitat in South Dakota and found no consistent trend in mass gain variation between birds utilizing either site type. In some cases, the Ovenbird gained significantly more mass in the anthropogenic woodlots than in the natural areas during fall migration stopover. One possible reason for this seemingly higher suitability of developed stopover areas compared to more natural areas is that urban areas can have ornamental vegetation, in addition to native vegetation, that provides more food diversity (in terms of berries and arthropods) for foragers (Liu & Swanson, 2014). The researchers also noted that while adult migrants are generally better at gaining mass during stopover, juveniles in their study sites were equally capable in this respect. Thus small, and even anthropogenic, forest fragments can serve as suitable stopover habitat for long-distance migrants. Our review indicates that many migrant species will seek out small forest fragments and use it as a stopover site. Historically, focus has been placed on conserving large areas of forest as this provides the most benefit to the greatest number of forest bird species. This review points out the importance of small forest patches, particularly to migrating birds in search of stopover sites. If preserving large forested areas is not feasible for a development scenario, planners can still attract a diverse array of migrants during spring and fall migration seasons if they preserve and incorporate tree canopy (including small forest patches, woodlots, and even individual trees in residential areas) into their landscape design. Planners’ goals to conserve forest fragments through development should be coupled with usage policies to be established post-development. For instance, if a developer decides to cluster houses away from forest fragments, strict usage restrictions that prohibit destructive activities in fragments (e.g. all-terrain vehicle use, off-leash pets, dumping, etc.), times limiting access, and so on, should be put in place to ensure the fragment is upkept (Farr, Pejchar, & Reed, 2017; Milder & Clark, 2011; Wald & Hostetler, 2010). These preserved forest fragments can also take the form of conservation easements—whereby landowners voluntarily cede land to government or land trust entities and receive a direct monetary or tax-reduction benefit in return—further incentivizing including them in development plans. Planning for postdevelopment should occur at the same time as planning for development and evaluated periodically to prevent deterioration of habitat.

built matrix, compared to adjacent forested areas, when native vegetation was incorporated in landscaping. All of this shows that city planners and developers that encourage the use of native vegetation can greatly improve the bird habitat in developed areas. Greater percentages of canopy cover in residential urban areas provide some vegetated complexity that more closely mimics natural forests (Belaire et al., 2014). In Arizona, a study by Green and Baker (2003) found that resident avian species richness was best predicted by canopy cover of woody species. Luther et al. (2008) found this to be particularly true for riparian-preferring species and Donnelly and Marzluff (2006) observed more native bird species in areas with more canopy cover in the Seattle, Washington metropolitan area. In addition, Hostetler and Holling (2000) found that birds’ numbers increased for each species with more tree canopy cover in cities. Additionally, snags are especially important to woodpecker species to forage for insects and to create cavities for nesting (Saab, Russell, & Dudley, 2009). Indeed, despite two studies (Parrish & Hepinstall-Cymerman, 2012; Schlesinger et al., 2008) from our Built Environment Review finding them to utilize residential areas during the breeding season, we maintain that the Pileated Woodpecker is an interior-forest specialist due to their reliance upon large forest habitat with snags. Snags should be left intact when safety to residents and homes is not compromised. Thus, land development regulations that stress the conservation of tree canopy and snags could greatly aid breeding and migrating species. 4.2.4. Urban dwellers Urban dwellers or synanthropic species are birds that successfully exploit anthropogenic changes and disturbances within an urban landscape while non-synanthropic species generally cannot (Fischer et al., 2015; Marzluff et al., 2012). Some urban dwellers are forest generalist species, such as the American Crow, that exploit the built environment for excess and discarded food waste created by humans in cities (Marzluff et al., 2012). Others are edge specialists, like the Brownheaded Cowbird, that take advantage of continuously disturbed habitat (Howell, Dijak, & Thompson, 2007). Others still, are almost completely integrated within the built matrix itself, trading tree cavity nests for cracks and crannies in buildings and other man-made structures (e.g. House sparrow and European starling) (Donnelly & Marzluff, 2006). Regardless of the type of habitat used, though, all urban dwellers increase in number where the spatial extent of urbanization increases. These species should always be expected to be common in built areas and in forests surrounded by residential areas. As such, it is important for planners to recognize that large populations of these urban dwelling species do not necessarily represent an urban design that benefits biodiversity. For instance, Donnelly and Marzluff (2006) found that increasing the amount of closed canopy, which generally increases the variety of bird species, actually reduced the gain of these urban dwellers. Planners should become familiar with urban dwelling species as indicators of ecological balance: if most of the city’s bird assemblage is made up of urban dwellers, planners can increase canopy cover and native vegetation in planned areas to increase the occurrence of nonurban dwellers. A simpler route to preserving natural bird diversity, though, would be to conserve forest fragments in developed areas. These conserved areas could take the form of many conventional features already implemented in urban planning today. Pocket parks that serve as both recreational areas as well as habitat for a variety woodland species; not only birds (Ikin et al., 2012). In some areas, fragmented forest patches can even be incorporated in stormwater management and design. Ecological buffer zones (e.g. forested riparian corridors) and setbacks (the regulatory policies that hold these zones in place) provide transitional areas that help mitigate harsh anthropogenic effects such as noise and lights from impacting natural areas. These tools empower planners to maintain complete tracts of forest (Schlacher, Weston, Lynn, & Connolly, 2013).

4.2.3. Built environment as habitat for forest birds Our review of forest birds observed in residential areas generated a list of 201 species. Indeed, a great many species of forest birds utilize built areas whether during the breeding or migration seasons. Importantly, for medium- and long-distance migrants, 46 species could use both forest fragments and residential areas as stopover sites during the spring and fall migration seasons, suggesting many migrants use residential landscapes with tree canopy cover as stopover sites. However, not all residential landscapes are created equal. Managing the quality of urban vegetation is key to creating good breeding and stopover habitat within the built environment and native vegetation can increase native bird diversity (Schneider & Miller, 2014). As found in one of our review studies, native bird density, diversity, richness, and biomass were all higher on residential properties landscaped with native vegetation (Burghardt et al., 2009). In Phoenix, Arizona, natural desert landscaping in urban neighborhoods increased the presence native desert avifauna and, together with proximity to larger desert tracts and higher-income neighborhoods, accounted for 50% of variation in diversity of the local urban bird community (Lerman & Warren, 2011). McCaffrey and Mannan (2012) showed that individual homeowners can increase the occurrence of native bird species based on small-scale landscaping and planting native vegetation in Tucson, Arizona. Mills et al. (1989) even found that native bird diversity was higher within the

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testing out an online platform for urban planners and developers: Building for Birds (http://wec.ifas.ufl.edu/buildingforbirds/web/home. html). This Web site offers users the opportunity to input general landscape data from current or future development projects and receive avian three separate habitat scores based on: a) the amount of forest fragments conserved as breeding habitat, b) the amount of different sizes of small, forest fragments that could be used as stopover habitat, and c) the amount of tree canopy conserved within residential areas. Combined with our avian species lists, the tool allows planners and developers to visualize which species are impacted during different seasons. This tool can be used to improve a development design to benefit certain groups of species. For example, if one wanted to improve the stopover habitat score for migrants, one could increase the sizes and density of relatively small forest fragments. This Building for Birds tool gives increasingly more points per acre when forest fragments sizes are increased. Further, in cases where forest fragment conservation is not feasible at all, points are still given for overall canopy cover in residential areas. Notably, these bird lists provide a snapshot and species presence does not imply thriving populations or persistence over time. We do not suggest that this species list conclusively states that these birds persist and survive just fine because other factors such as disease, competition, and predators may negatively affect certain species. However, planners can use this tool to first identify local species of concern and then collaborate with ecologists to assess population vitality and conduct further research to see, for example, whether species listed breed successfully and/or find enough forage in these fragmented and residential areas during migration. With this, both parties have a seat at the table and can work together to better devise and implement appropriate development strategies for managing meaningful bird habitat.

4.3. Beyond design While it is important to conserve forest fragments as development and urbanization increase in neighborhoods, taking construction considerations and post-construction management of these fragments into account is a major factor in the success of biodiversity conservation in the built environment (Hostetler, 2012). Despite employing “green” design in their structures, construction practices such as parking heavy machinery in natural areas, failure to protect root zones with proper fencing, and failure to recognize and remove invasive vegetation transported from other areas can greatly reduce soil quality and, ultimately, reduce ecosystem integrity. Setting aside natural, undeveloped areas within an urban area has very little positive effects for biodiversity when steps are not taken to ensure its biological integrity. Without long-term management, forest fragments can become ecological traps—habitat that an organism might favor despite increased likelihood of species mortality and decline (Battin, 2004). For example, Shipley, Murphy, and Elzinga (2013) found that Spotted Towhees nesting on the edges of urban forest experienced low post-fledging success because of increased predation by free-ranging pets (e.g. cats and dogs). The introduction of exotic plants can lead to invasions of forest fragments and landscape homogenization, a fundamental weakening of biological integrity that gives way to further invasions and even local extinctions (Schneider & Miller, 2014). In forest fragments, executing prescribed burn and herbicidal treatment strategies can aid in weeding out invasive plants and maintain native plant diversity (Huston, 2004; Provencher et al., 2001), but this is difficult to do in urban areas without citizen acceptance. The introduction of direct human disturbance within conserved forest fragments should also be given substantial management concern. Off-road vehicle use, noise pollution, and waste dumping are only a few examples of habitat-destructive practices that often accompany urban development (e.g., Sauvajot, Buechner, Kamradt, & Schonewald, 1998). Even seemingly innocuous recreational trails can dramatically change the density and behavior of local species (Bhardwaj, Dale, & Ratcliffe, 2015). Managing (and reducing) these impacts on conserved areas within or adjacent to the built matrix may be as simple as erecting prohibitive signage or implementing educational programs to educate residents (Hostetler & Drake, 2009; Hostetler, Swiman, Prizzia, & Noiseux, 2008). Without post-development planning and management, conserved areas will not function optimally for species that inhabit them in the long term. Equally important, several studies have shown that with higher bird diversity comes greater public appreciation of private and public property (Hedblom, Heyman, Antonsson, & Gunnarsson, 2014). This parallels a growing body of research that shows healthy natural environments increase human well-being and community engagement (Soulsbury & White, 2015). As the link has clearly been drawn between conservation of forest fragments and increased bird biodiversity (Green & Baker, 2003), planners and developers who conserve forest fragments in and around the built matrix help communities gain both ecological and social benefits.

5. Conclusions The strength of this review lies in its empirical evidence comes from both urban and rural fragmentation studies. This systematic review revealed that forest fragments and residential areas with trees in urban areas can be used by a variety of species and should not be discounted when cities are trying to conserve bird diversity. For city planners and developers alike, it takes considerable effort to conserve individual trees and small forest fragments when land is subdivided. This means that adequate fencing needs to be installed to protect trees, roads and buildings be aligned to maximize conservation of small forest fragments, and built areas are designed with native vegetation and plans to protect tree canopy cover. Thus, even when large forest conservation is not an option for a site, a host of built environment professionals and ecologists can still collaborate and maximize the conservation of avian habitat. Conserving forest fragments and individual trees can be incorporated into a site design as pocket parks, stormwater retention areas, and buffer strips. The bird lists generated in this review revealed that efforts to conserve even individual trees pays dividends for a variety of species, including some interior-forest specialists and endangered species. Forest fragments serving as bird habitat can be found at all scales of urban planning from conserving trees along a shared property line, to designing pocket parks within neighborhoods, to setting aside large conservation areas at the municipal level. In the end, to conserve bird habitat, thoughtful planning and construction management is needed, and this review highlights the importance of conserving individual trees in residential areas and even the smallest of forest fragments.

4.4. Use of bird lists in planning The generated bird lists can even be used to develop evaluation tools where different development scenarios would output different avian habitat scores. When landscape fragmentation is already inherent within a proposed development site, site designs can be evaluated for avian habitat across migration and breeding seasons. We currently are Appendix A Table A1

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Archilochus colubris Selasphorus sasin Calypte anna Archilochus alexandri Cynanthus latirostris Selasphorus calliope Calypte costae Selasphorus rufus Columbina passerina Streptopelia decaocto Zenaida macroura Columbina inca Patagioenas fasciata Zenaida asiatica Coccyzus erythropthalmus Coccyzus americanus Empidonax virescens Icterus galbula Dendroica castanea Mniotilta varia Dendroica fusca Dendroica caerulescens Dendroica virens Vireo altiloquus Cyanocitta cristata Vireo solitarius Vermivora cyanoptera Quiscalus major Toxostoma rufum Sitta pusilla Wilsonia canadensis Parus carolinensis Thryothorus ludovicianus Dendroica cerulea Dendroica pensylvanica Oporornis agilis Spiza americana Sialia sialis Sayornis phoebe Pipilo erythrophthalmus Contopus virens Spizella pusilla Corvus ossifragus Vermivora chrysoptera Dumetella carolinensis

Ruby-throated Hummingbird Allen's Hummingbird Anna's Hummingbird Black-chinned Hummingbird Broad-billed Hummingbird Calliope Hummingbird Costa's Hummingbird Rufous Hummingbird Common Ground-dove Eurasian Collared-dove Mourning Dove Inca Dove Northern Band-tailed Pigeon White-winged Dove Black-billed Cuckoo Yellow-billed Cuckoo Acadian Flycatcher Baltimore Oriole Bay-breasted Warbler Black-and-white Warbler Blackburnian Warbler Black-throated Blue Warbler Black-throated Green Warbler Black-whiskered Vireo Blue Jay Blue-headed Vireo Blue-winged Warbler Boat-tailed Grackle Brown Thrasher Brown-headed Nuthatch Canada Warbler Carolina Chickadee Carolina Wren Cerulean Warbler Chestnut-sided Warbler Connecticut Warbler Dickcissel Eastern Bluebird Eastern Phoebe Eastern Towhee Eastern Wood-pewee Field Sparrow Fish Crow Golden-winged Warbler Grey Catbird

Apodiformes Apodiformes Apodiformes Apodiformes Apodiformes Apodiformes Apodiformes Apodiformes Columbiformes Columbiformes Columbiformes Columbiformes Columbiformes Columbiformes Cuculiformes Cuculiformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes

E W W W W W W W E/ W E/ W E/ W W W W E E E E E E E E E E E E E E E E E E E E E E E E E E E E E E E

LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC VU LC LC LC LC LC LC LC LC LC NT LC

IUCN CODE2

Common Name

Order

Locality

1

Scientific Name

Natural History

Species

OW OW OW OW SC OW D OW SC T OW T F OW F OW F OW F F F F F F F F OW MSW SC F F F OW F OW F FE FE OW SC F SC SL OW OW

Habitat Type3 MD/ LD LD YR MD/ LD SD LD YR/ SD LD YR/ SD YR YR/ LD YR YR/ MD YR/ SD LD LD LD MD/ LD LD YR/ SD/ LD LD LD LD LD YR/ SD LD LD YR YR/ SD YR LD YR YR LD LD LD LD YR/ MD YR/ SD/ MD YR/ SD LD YR/ MD YR/ SD LD YR/ MD

Migrant Status4 yes – – – – – – – – – yes – – – yes yes yes yes – yes – yes yes – yes – no – no – yes yes yes yes yes yes – – – yes yes – yes – yes

Breeds in LateSuccessional Forest5

Breeding Review

yes – – – – – – – – – yes – – – no yes no yes – no – no no – yes – yes – no – no yes yes no yes no – – – yes yes – yes – yes

Breeds in EarlySuccessional Forest6 (1/2) – – – – – – – – – (0/2) – – – (1/2) Low (1/4) High (6/6) Low (1/3) – High (3/3) – (2/2) (1/2) – No (0/3) – (0/1) – (1/1) – (2/2) (0/1) (1/2) High (4/4) (1/2) (1/1) – – – (0/2) No (0/5) – (0/1) – (0/2)

Confidence in InteriorForest Specialist Assignment7 2 – – 1 – – 1 – 1 – 3 – – 1 1 2 1 3 4 5 7 9 6 1 4 7 3 – 1 – 6 – – 1 8 2 – – 5 2 7 2 1 3 8

(continued on next page)

Total no. of Studies That Observed Species in Small Forest Fragments during Migration Seasons

Stopover Review

Table A1 COMBINED AVIAN SPECIES LIST FROM ALL THREE SYSTEMATIC REVIEWS. Combined Avian Species List from All Three Reviews. This table combines the species lists generated by each systematic review in our study. The birds in this list are those species that are tree-nesting species or species associated with trees. The Breeding Review columns show which species will breed in late or early successional forest fragments as well as which species are Interior-Forest Specialists (birds that do not breed in forest fragments). The Stopover Review column lists which species were observed small forest fragments by studies conducted during the spring and fall migration seasons. The Built Environment Review columns show which species were observed within residential areas and gives the season of the observation. The Synanthropic Review columns show which species are synanthropic (urban-adapted species commonly found within the built matrix). Species are listed alphabetically by taxa and common name.

J.-M.J. Archer et al.

Landscape and Urban Planning 185 (2019) 1–23

11

Tyrannus dominicensis Catharus minimus Myiarchus crinitus Icterus cucullatus Wilsonia citrina Passerina cyanea Oporornis formosus Parkesia motacilla Oporornis philadelphia Vermivora ruficapilla Cardinalis cardinalis Parula americana Icterus spurius Vireo philadelphicus Dendroica pinus Dendroica discolor Protonotaria citrea Pheucticus ludovicianus Piranga olivacea Limnothlypis swainsonii Baeolophus bicolor Vireo griseus Hylocichla mustelina Helmitheros vermivorum Empidonax flaviventris Vireo flavifrons Dendroica dominica Corvus brachyrhynchos Carduelis tristis Setophaga ruticilla Turdus migratorius Certhia americana Hirundo rustica Vireo bellii Thryomanes bewickii Parus atricapillus Dendroica striata Passerina caerulea Polioptila caerulea Euphagus cyanocephalus Molothrus ater Dendroica tigrina Bombycilla cedrorum Spizella passerina Spizella pallida Quiscalus quiscula Corvus corax Sturnus vulgaris Geothlypis trichas Toxostoma curvirostre Junco hyemalis

Grey Kingbird Grey-cheeked Thrush Great Crested Flycatcher Hooded Oriole Hooded Warbler Indigo Bunting Kentucky Warbler Louisiana Waterthrush Mourning Warbler Nashville Warbler Northern Cardinal Northern Parula Orchard Oriole Philadelphia Vireo Pine Warbler Prairie Warbler Prothonotary Warbler Rose-breasted Grosbeak Scarlet Tanager Swainson's Warbler Tufted Titmouse White-eyed Vireo Wood Thrush Worm-eating Warbler Yellow-bellied Flycatcher Yellow-throated Vireo Yellow-throated Warbler American Crow American Goldfinch American Redstart American Robin American Treecreeper Barn Swallow Bell's Vireo Bewick's Wren Black-capped Chickadee Blackpoll Warbler Blue Grosbeak Blue-grey Gnatcatcher Brewer's Blackbird Brown-headed Cowbird Cape May Warbler Cedar Waxwing Chipping Sparrow Clay-coloured Sparrow Common Grackle Common Raven Common Starling Common Yellowthroat Curve-billed Thrasher Dark-eyed Junco

Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes

E E E E E E E E E E E E E E E E E E E E E E E E E E E E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ E/ W W W W W W W W W W W W W W W W W W W W W W W W

LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC NT LC LC LC LC LC LC LC LC LC LC NT LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC

IUCN CODE2

Common Name

Order

Locality

1

Scientific Name

Natural History

Species

Table A1 (continued)

OW F OW F F OW F RF F F OW F OW F F OW F F F F F SC F F F OW F OW OW F OW F T SC OW F F OW F T FE F OW OW SC OW MTW T SC SC F

Habitat Type3 LD LD MD/ LD LD LD LD LD LD LD LD YR LD LD LD MD MD LD LD LD MD YR MD LD LD YR/ SD/ LD LD LD YR/ SD YR/ SD LD YR/ SD YR/ SD LD MD YR YR/ IR LD LD YR/ LD YR/ MD YR/ SD LD YR/ SD/ LD YR/ SD MD/ LD YR/ SD YR YR/ SD YR/ LD YR YR/ MD

Migrant Status4 – – yes – yes yes yes yes yes – yes yes – – yes yes yes yes yes – yes yes yes yes – yes yes yes no yes yes yes – – – yes – – yes – yes – yes no – yes yes yes no – –

Breeds in LateSuccessional Forest5

Breeding Review

– – yes – no yes no no no – yes no – – – yes no no no – yes yes no no – no no yes yes no yes no – – – yes – – no – yes – yes no – yes yes yes yes – –

Breeds in EarlySuccessional Forest6 – – Low (1/7) – Med (2/3) No (0/4) High (3/4) (2/2) (1/1) – No (0/5) High (4/4) – – (1/2) (0/1) (0/1) High (3/4) High (7/7) – Low (1/5) (0/2) Med (5/8) (2/2) – High (3/4) (1/2) Low (1/3) (0/2) Med (2/3) No (0/5) (0/1) – – – No (0/3) – – High (4/4) – No (0/3) – (0/2) (0/1) – No (0/3) (0/1) No (0/4) (0/2) – –

Confidence in InteriorForest Specialist Assignment7 1 5 6 2 1 7 – 4 6 10 – 9 2 5 5 3 2 7 8 1 – 4 7 3 5 2 2 – 3 12 2 4 – 1 – 3 11 1 7 – 5 4 5 2 1 2 – 1 8 – 2

(continued on next page)

Total no. of Studies That Observed Species in Small Forest Fragments during Migration Seasons

Stopover Review

J.-M.J. Archer et al.

Landscape and Urban Planning 185 (2019) 1–23

12

Tyrannus tyrannus Coccothraustes vespertinus Passerella iliaca Regulus satrapa Zonotrichia querula Catharus guttatus Carpodacus mexicanus Passer domesticus Troglodytes aedon Chondestes grammacus Empidonax minimus Melospiza lincolnii Lanius ludovicianus Dendroica magnolia Cistothorus palustris Mimus polyglottos Parkesia noveboracensis Contopus cooperi Vermivora celata Seiurus aurocapilla Dendroica palmarum Pinicola enucleator Carduelis pinus Carpodacus purpureus Progne subis Cardinalis sinuatus Loxia curvirostra Sitta canadensis Vireo olivaceus Agelaius phoeniceus Regulus calendula Melospiza melodia Piranga rubra Catharus ustulatus Melospiza georgiana Vermivora peregrina Tachycineta bicolor Catharus fuscescens Vireo gilvus Sitta carolinensis Zonotrichia leucophrys Zonotrichia albicollis Empidonax traillii Wilsonia pusilla Troglodytes troglodytes Dendroica petechia Icteria virens Dendroica coronata Perisoreus canadensis Passerina ciris Melozone aberti

Eastern Kingbird Evening Grosbeak Fox Sparrow Golden-crowned Kinglet Harris's Sparrow Hermit Thrush House Finch House Sparrow House Wren Lark Sparrow Least Flycatcher Lincoln's Sparrow Loggerhead Shrike Magnolia Warbler Marsh Wren Northern Mockingbird Northern Waterthrush Olive-sided Flycatcher Orange-crowned Warbler Ovenbird Palm Warbler Pine Grosbeak Pine Siskin Purple Finch Purple Martin Pyrrhuloxia Red Crossbill Red-breasted Nuthatch Red-eyed Vireo Red-winged Blackbird Ruby-crowned Kinglet Song Sparrow Summer Tanager Swainson's Thrush Swamp Sparrow Tennessee Warbler Tree Swallow Veery Warbling Vireo White-breasted Nuthatch White-crowned Sparrow White-throated Sparrow Willow Flycatcher Wilson's Warbler Winter Wren Yellow Warbler Yellow-breasted Chat Yellow-rumped Warbler Grey Jay Painted Bunting Abert's Towhee

Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes

E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/ W E/W E/W W

LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC NT LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC NT LC

IUCN CODE2

Common Name

Order

Locality

1

Scientific Name

Natural History

Species

Table A1 (continued)

FE F F F F OW T T OW FE F SC OW F MSW T F OW F F OW OW OW F LPF/T SC F F F MSW F OW OW F MSW F LPF F OW F SC F MSW SC F OW SC F F SC D

Habitat Type3 LD YR/IR YR/ SD/ YR/ MD MD YR/ SD YR/ SD YR YR/ SD/ MD LD LD YR/MD LD YR/MD YR LD LD MD/ LD LD LD YR YR/IR YR/ SD LD YR MD YR/ SD/ LD YR/ SD YR/ SD YR/ MD LD LD LD LD LD LD MD/ LD YR YR/ MD SD/ MD LD LD LD LD LD YR/ SD/ YR MD YR

Migrant Status4

13 LD

IR

MD

LD

– – – – – – – – yes – yes – – yes – – yes – – yes – – – – – – – – yes yes – yes yes no – – – yes – yes – yes – – – – yes – – – –

Breeds in LateSuccessional Forest5

Breeding Review

– – – – – – – – yes – yes – – no – – no – – no – – – – – – – – no yes – yes no no – – – no – no – yes – – – – yes – – – –

Breeds in EarlySuccessional Forest6 – – – – – – – – (0/2) – (1/1) – – (0/1) – – High (3/3) – – High (6/6) – – – – – – – – Med (4/7) No (0/4) – (0/1) (2/2) (1/1) – – – High (4/4) – Med (2/3) – (0/1) – – – – (0/2) – (0/1) – –

Confidence in InteriorForest Specialist Assignment7 – 1 1 4 1 3 – – 3 – 6 4 – 12 – – 8 2 4 11 5 – 2 3 – – – 3 11 2 8 2 2 10 3 7 1 5 4 – 1 3 5 8 2 7 3 11 – 2 –

(continued on next page)

Total no. of Studies That Observed Species in Small Forest Fragments during Migration Seasons

Stopover Review

J.-M.J. Archer et al.

Landscape and Urban Planning 185 (2019) 1–23

California Thrasher California Towhee Canyon Towhee Cassin's Finch Cassin's Kingbird Cassin's Vireo Chestnut-backed Chickadee Clark's Nutcracker Great-tailed Grackle Green-tailed Towhee Grey Flycatcher Hammond's Flycatcher Hermit Warbler Hutton's Vireo Lazuli Bunting Le Conte's Thrasher Lesser Goldfinch Lucy's Warbler MacGillivray's Warbler Mountain Chickadee North-western Crow Oak Titmouse Pacific-slope Flycatcher Phainopepla Pygmy Nuthatch Rufous-crowned Sparrow Rufous-winged Sparrow Sage Thrasher Sagebrush Sparrow Spotted Towhee Steller's Jay Townsend's Solitaire Townsend's Warbler Tropical Kingbird Varied Thrush Verdin Violet-green Swallow

Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes

Empidonax oberholseri Myiarchus cinerascens Toxostoma bendirei Sayornis nigricans Pheucticus melanocephalus Polioptila melanura Dendroica nigrescens Amphispiza bilineata Spizella breweri Molothrus aeneus Icterus bullockii Psaltriparus minimus Campylorhynchus brunneicapillus Toxostoma redivivum Melozone crissalis Melozone fuscus Carpodacus cassinii Tyrannus vociferans Vireo cassinii Parus rufescens Nucifraga columbiana Quiscalus mexicanus Pipilo chlorurus Empidonax wrightii Empidonax hammondii Dendroica occidentalis Vireo huttoni Passerina amoena Toxostoma lecontei Carduelis psaltria Vermivora luciae Oporornis tolmiei Parus gambeli Corvus caurinus Baeolophus inornatus Empidonax difficilis Phainopepla nitens Sitta pygmaea Aimophila ruficeps Peucaea carpalis Oreoscoptes montanus Artemisiospiza nevadensis Pipilo maculatus Cyanocitta stelleri Myadestes townsendi Dendroica townsendi Tyrannus melancholicus Zoothera naevia Auriparus flaviceps Tachycineta thalassina

American Dusky Flycatcher Ash-throated Flycatcher Bendire's Thrasher Black Phoebe Black-headed Grosbeak Black-tailed Gnatcatcher Black-throated Grey Warbler Black-throated Sparrow Brewer's Sparrow Bronzed Cowbird Bullock's Oriole Bushtit Cactus Wren

Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes W W W W W W W W W W W W W W W W W W W W W W W W W W W W W W W W W W W W W

W W W W W W W W W W W W W LC LC LC NT LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC

LC LC VU LC LC LC LC LC LC LC LC LC LC

IUCN CODE2

Common Name

Order

Locality

1

Scientific Name

Natural History

Species

Table A1 (continued)

SC SC SC F OW F F F T SC OW F F F OW SC OW OW OW F OW OW F SC F SC SC SC SC SC F OW F OW F SC OW

OW OW D OW F SC F SC SC T OW SC D

Habitat Type3

YR YR YR YR/ MD LD YR YR/ YR YR/ LD LD LD YR LD YR YR/ MD LD YR YR YR LD MD YR YR YR YR/ YR/ YR/ YR LD LD LD YR/ YR LD

14 SD

SD SD SD

SD

SD/ MD

SD

MD

LD LD YR/ SD YR/ SD MD/ LD YR LD LD YR/ SD LD MD YR YR

Migrant Status4

– – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – –

– – – yes – – – – – – – – –

Breeds in LateSuccessional Forest5

Breeding Review

– – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – –

– – – yes – – – – – – – – –

Breeds in EarlySuccessional Forest6

– – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – –

– – – (0/1) – – – – – – – – –

Confidence in InteriorForest Specialist Assignment7

– – – – – – – – – – 1 – 1 – – – – – – – – – 1 2 – – – – – – – – – – – – –

– 1 – – 1 – – – – – 1 – –

(continued on next page)

Total no. of Studies That Observed Species in Small Forest Fragments during Migration Seasons

Stopover Review

J.-M.J. Archer et al.

Landscape and Urban Planning 185 (2019) 1–23

Built Environment Review

No. of Studies That Observed Species in the Built Environment during the Breeding Season

3 1 4 2 1 1 1 3 1 2 8 2 3 3 1 2 2 5 0 2 – 1

Species

Order

15

Apodiformes Apodiformes Apodiformes Apodiformes Apodiformes Apodiformes Apodiformes Apodiformes Columbiformes Columbiformes Columbiformes Columbiformes Columbiformes Columbiformes Cuculiformes Cuculiformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes

W W W W W W W E E E/ E/ E/ E/ E/ W W W W W W W W W W W W W

1 0 0 0 0 0 0 1 0 0 0 0 0 0 1 2 1 2 1 1 – 1

No. of Studies That Observed Species in the Built Environment during the Spring or Fall Migration Seasons

Sialia mexicana Tyrannus verticalis Aphelocoma californica Piranga ludoviciana Contopus sordidulus Loxia leucoptera Chamaea fasciata Melanerpes carolinus Melanerpes erythrocephalus Dryobates pubescens Leuconotopicus villosus Hylatomus pileatus Sphyrapicus varius Colaptes auratus Melanerpes formicivorus Melanerpes uropygialis Colaptes chrysoides Dryobates scalaris Dryobates nuttallii Sphyrapicus ruber Leuconotopicus albolarvatus Sphyrapicus thyroideus

Western Bluebird Western Kingbird Western Scrub-jay Western Tanager Western Wood-pewee White-winged Crossbill Wrentit Red-bellied Woodpecker Red-headed Woodpecker Downy Woodpecker Hairy Woodpecker Pileated Woodpecker Yellow-bellied Sapsucker Yellow-shafted Flicker Acorn Woodpecker Gila Woodpecker Gilded Flicker Ladder-backed Woodpecker Nuttall's Woodpecker Red-breasted Sapsucker White-headed Woodpecker Williamson's Sapsucker

Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes

OW FE SC F OW OW SC F OW F F F F OW OW D D D OW F F F

Habitat Type3

0 0 3 3 0 0 2 1 1 0 3 3 0 2 0 0 0 1 0 0 – 0

No. of Studies that Observed Species (Built Env.) during Breed-Migration8

LC LC LC LC LC LC LC LC NT LC LC LC LC LC LC LC LC LC LC LC LC LC

IUCN CODE2

Common Name

Order

Locality

1

Scientific Name

Natural History

Species

Table A1 (continued)

– – yes – – – – yes yes yes yes yes yes yes yes – – – – – – –

Breeds in LateSuccessional Forest5

(1/1) – High (3/3) (1/1) – – – (2/2) – – High (3/3) – (2/2) – – – (0/1) – – (0/1) – –

Confidence in Synanthropic Species Assignment9

Synanthropy Analysis

YR/MD MD/ LD YR LD LD YR/ SD YR YR YR/ SD YR YR YR MD/ LD YR/ SD YR YR YR YR YR MD YR YR/ SD

Migrant Status4

Breeding Review

(0/1) – (0/1) – – – – Med (3/5) (0/1) No (0/3) High (4/4) High (3/3) (0/1) (0/1) (0/1) – – – – – – –

Confidence in InteriorForest Specialist Assignment7

?

X

? ✓* ✓ ✓ ✓* ? ✓ X ✓

? ✓

✓ ? ?

✓*

Breeds in Forest Fragment

✓ ✓* ? ✓ ✓ ✓ ✓ ✓ ✓ ✓

✓

✓* ? ? ? ✓ ?

✓ ?

Stopover in Forest Fragment

Overall Comparisons

– – yes – – – – no no no no no no yes yes – – – – – – –

Breeds in EarlySuccessional Forest6

X*

✓ ✓ ? ? ✓ ✓ ✓ ✓ ? ✓ X* ?

✓ ? ?

✓

?

✓ ✓* ? ✓ ✓ ✓ ? ✓

✓

✓ ? ? ? ✓ ?

✓ ?

Stopover in Residential Area

(continued on next page)

Breeds in Residential Area

– 1 – – – 1 – – 1 – – – 2 4 – – – – – – – –

Total no. of Studies That Observed Species in Small Forest Fragments during Migration Seasons

Stopover Review

J.-M.J. Archer et al.

Landscape and Urban Planning 185 (2019) 1–23

Built Environment Review

No. of Studies That Observed Species in the Built Environment during the Breeding Season

1 – 5 0 1 0 5 1 0 4 3 – 0 0 1 4 5 4 6 2 – – 6 – – 4 1 2 5 1 0 – 1 9 1 0 – 1 1 0 2 5 1 4 1 5 0 0 2 0 7

Species

Order

Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes

Table A1 (continued)

1 – 1 2 1 0 1 0 2 0 1 – 1 0 0 0 0 0 1 0 – – 2 – – 2 0 1 2 2 1 – 0 1 1 1 – 0 1 1 2 1 0 1 2 1 1 1 1 1 1

No. of Studies That Observed Species in the Built Environment during the Spring or Fall Migration Seasons 0 – 0 1 0 1 0 0 0 0 0 – 0 1 0 0 0 1 0 0 – – 0 – – 0 0 1 0 0 0 – 1 2 0 0 – 0 0 0 0 0 0 0 0 0 0 0 0 0 1

No. of Studies that Observed Species (Built Env.) during Breed-Migration8 – – (1/1) – – – (0/1) (0/1) – (1/1) (0/1) – – – – (1/1) (1/1) (0/1) (0/1) (1/2) – – (1/1) – – (0/1) – (0/1) No (0/1) – – – (0/1) (1/1) (0/1) – – (0/1) (0/1) – – (0/1) – (1/1) (0/1) (0/1) – – (0/1) – High (4/4)

Confidence in Synanthropic Species Assignment9

Synanthropy Analysis

16

X ? ✓

✓ ? X ?

? ✓* ? X X

✓ X

X ✓ X ? ?

✓

✓

?

✓ ✓* ✓* ✓

? ? ✓ ? ? ✓ ✓ X ? ?

✓

?

Breeds in Forest Fragment

✓ ✓ ✓ ✓ ✓ ✓ ✓

✓ ? ✓ ✓ ✓ ? ✓ ✓ ?

? ✓ ? ? ✓ ✓ ✓ ✓ ? ? ✓ ✓ ? ✓ ✓ ? ? ✓ ✓* ✓ ✓ ✓

✓

✓*

✓ ? ✓ ✓ ✓

Stopover in Forest Fragment

Overall Comparisons

✓

X*

X*

✓

X* X*

?

✓ X*

✓ ? X* ✓ X*

✓

✓ ✓ ✓ ✓ ?

✓ ✓ X*

? ✓ ?

✓

✓ ✓ ? ? ✓ ? ✓*

? ✓ ? ✓* ✓ ?

? ?

?

✓* ✓ ?

✓

✓

? ? ? ✓ ✓* ✓* ✓* ?

?

✓*

✓

✓

?

Stopover in Residential Area

(continued on next page)

Breeds in Residential Area

J.-M.J. Archer et al.

Landscape and Urban Planning 185 (2019) 1–23

Built Environment Review

No. of Studies That Observed Species in the Built Environment during the Breeding Season

7 2 8 4 4 1 2 5 2 2 3 1 9 0 6 7 – 5 1 8 3 4 2 4 1 1 3 – 2 10 9 6 0 1 0 1 0 1 6 0 3 3 3 0 1 3 3 1 3 2 3

Species

Order

Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes

Table A1 (continued)

1 2 0 0 0 0 1 1 2 1 2 0 0 1 1 0 – 0 0 0 1 0 0 2 0 0 1 – 1 0 0 0 0 1 0 0 2 0 1 1 1 1 2 1 0 1 0 0 0 1 1

No. of Studies That Observed Species in the Built Environment during the Spring or Fall Migration Seasons 1 0 2 0 1 0 3 0 0 0 2 1 3 0 0 1 – 0 3 2 2 2 2 0 0 0 0 – 1 3 1 2 2 0 2 1 0 1 3 0 0 3 0 0 0 0 0 0 1 0 1

No. of Studies that Observed Species (Built Env.) during Breed-Migration8 Med (2/3) – Low (1/3) (0/2) (1/1) – Med (2/3) (2/2) – (0/1) (0/1) – High (4/4) – (2/2) – – (1/1) (1/1) High (4/4) (1/2) – (0/2) – (1/1) – (0/1) – – High (4/4) (1/1) (2/2) – – – – – – (2/2) – (2/2) (2/2) – – (1/1) (1/1) (2/2) – – (1/1) (0/2)

Confidence in Synanthropic Species Assignment9

Synanthropy Analysis

17

✓*

?

? ✓* ✓*

✓ X ✓* ✓* X

? ?

✓

✓ ✓ ✓*

✓*

✓ ✓ ✓ ✓ ✓ ?

✓* ✓

X ? ✓

✓ X ✓* ✓* ✓ ? ✓ ✓

Breeds in Forest Fragment

✓ ✓

✓ ✓ ?

✓ ✓ ✓ ✓ ✓

✓ ? ✓ ✓ ? ✓ ?

✓ ✓ ? ? ✓ ? ✓ ✓

✓* ✓

✓ ✓ ✓ ? ✓ ✓ ✓ ✓ ? ✓*

✓ ✓ ✓* ✓ ✓* ?

Stopover in Forest Fragment

Overall Comparisons

✓

?

? ✓ ✓

✓ X* ✓ ✓ X*

?

✓ ✓ ✓

✓

✓ ✓ ✓ ✓ ✓ ?

✓ ✓

X* ? ✓

✓ X* ✓ ✓ ✓ ? ✓ ✓

✓ ✓*

✓ ✓* ?

? ✓ ✓ ✓ ?

✓ ? ? ? ? ? ?

✓ ✓

? ✓*

✓ ✓

✓ ✓

✓*

✓ ✓ ✓ ? ✓ ? ✓* ?

✓ ✓ ✓ ✓* ✓* ?

Stopover in Residential Area

(continued on next page)

Breeds in Residential Area

J.-M.J. Archer et al.

Landscape and Urban Planning 185 (2019) 1–23

Built Environment Review

No. of Studies That Observed Species in the Built Environment during the Breeding Season

4 7 0 8 2 – 0 1 5 2 5 7 0 – 4 3 1 6 0 2 – 0 1 1 3 2 – 5 3 2 3 1 2 1 2 4 0 0 1 1 1 3 1 1 3 1 0 1 1 2 0

Species

Order

Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes

Table A1 (continued)

2 0 1 0 2 – 1 1 1 2 0 1 0 – 1 1 1 2 1 0 – 0 0 0 0 0 – 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0

No. of Studies That Observed Species in the Built Environment during the Spring or Fall Migration Seasons 0 2 2 2 0 – 0 0 0 0 0 1 2 – 0 2 0 2 1 1 – 0 2 1 0 1 1 2 2 1 2 2 2 1 2 3 1 1 2 0 0 0 0 0 1 1 1 1 1 1 1

No. of Studies that Observed Species (Built Env.) during Breed-Migration8 (1/1) (2/2) – High (3/4) No (0/1) No (0/2) – – – – (2/2) Med (2/3) (1/1) – (1/1) No (0/3) (0/2) (1/2) – – (1/1) – – – (0/2) – (2/2) Med (2/3) – No (0/2) – – – (1/2) High (3/3) – (0/1) (1/1) – – – (2/2) (0/2) – – – – (0/1) (0/1) No (0/3) (1/1)

Confidence in Synanthropic Species Assignment9

Synanthropy Analysis

18

?

✓ ? ? ✓

✓ ✓ ? ? ✓

?

? ✓ ✓ ✓

✓

?

?

?

✓

X

✓ ✓ ✓ ✓ ? ?

Breeds in Forest Fragment

?

✓ ? ? ?

?

? ? ✓

✓* ✓* ✓ ✓ ?

? ✓ ? ✓ ✓

?

✓ ✓ ✓ ✓ ? ✓ ✓ ✓

✓ ✓ ✓ ✓* ✓ ✓ ✓ ✓ ✓* ✓ ✓

Stopover in Forest Fragment

Overall Comparisons

?

✓ ? ? ✓

✓ ✓ ? ? ✓

?

? ? ✓ ✓

✓

?

✓

X*

✓ ✓ ? ✓

? ? ? ?

?

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?

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Stopover in Residential Area

(continued on next page)

Breeds in Residential Area

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Landscape and Urban Planning 185 (2019) 1–23

19

No. of Studies That Observed Species in the Built Environment during the Breeding Season

1 3 1 3 1 1 0 1 2 1 0 2 0 0 2 1 1 1 0 0 4 1 0 2 0 3 3 – 0 6 3 8 7 2 0 10 0 4 0 2 0 1 1 1

Order

Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Passeriformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes Piciformes

0 0 0 1 0 1 0 0 0 0 0 0 0 0 1 1 1 0 0 1 0 0 0 0 0 1 1 – 0 1 1 2 1 0 1 1 0 0 0 0 0 0 0 0

No. of Studies That Observed Species in the Built Environment during the Spring or Fall Migration Seasons 0 3 2 1 0 0 1 1 3 0 1 0 1 1 0 0 0 1 1 0 2 1 0 2 1 1 1 – 1 0 0 1 0 0 1 1 1 2 1 2 1 0 0 0

No. of Studies that Observed Species (Built Env.) during Breed-Migration8 – (1/2) – (2/2) – – (0/1) No (0/3) – – – – – – (0/3) (0/2) – (0/1) – – – (2/2) (0/1) No (0/1) (2/2) (0/2) (2/2) – – (1/1) (0/1) Low (1/4) Low (1/3) (0/2) – High (3/4) High (4/4) – – – (0/1) (0/1) – –

Confidence in Synanthropic Species Assignment9

Synanthropy Analysis

? ?

✓* ✓ ✓ ? ✓ ?

? ✓ ✓* ✓ ✓ X

?

✓

✓* ? ? ? ? ? ? ?

✓ ? ? ?

? ✓

Breeds in Forest Fragment

?

✓ ✓

✓*

✓ ✓ ?

? ? ✓*

? ? ? ?

? ? ?

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Stopover in Forest Fragment

Overall Comparisons

? ?

✓ ✓ ✓ ? ✓ ?

? ✓ ✓ ✓ ✓ X*

?

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✓ ? ? ? ? ? ? ?

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Breeds in Residential Area

?

? ✓*

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Stopover in Residential Area

2

Locality (Regional Occurrence): East (E) = Occurs east of the 100th meridian, West (W) = occurs west of the 100th meridian. The IUCN (International Union for Conservation of Nature) identifies the conservation status of species all over the world. Least Concern (LC) represents species that have the lowest risk of becoming endangered in the wild. Near Threatened (NT) represents species that are likely to become threatened in the near future (e.g. due to increasing trends in habitat loss). Vulnerable (VU) represents species that are at high risk of becoming endangered in the wild (due to current and ongoing threats). 3 Habitat Types: Forest (F) = mature, forest fragments; Open woodland (OW) = disturbed or regrowing forest; Scrub (SC) = dense shrubbery, including abandoned farm fields, clearcuts, powerline corridors, fencerows, forest edges and openings, swamps, and edges of streams and ponds; Marsh woods (MSW) = various wetlands, including freshwater and tidal marshes, bogs, meadows, and swamps; Mountain Woods

1

Built Environment Review

Species

Table A1 (continued)

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(MTW) = mountain forests; Forest edge (FE) = disturbed habitat, similar to early successional forest, at the edge of a forest; Riparian forest (RF) = forest buffer along a river or waterway; Lake/Pond Forest (LPF) = forest surrounding a body of water; [Allaboutbirds.org, Cornell Bird Lab]. Note that if a species is not an interior forest specialist and it breeds in mature forest, that means it would either breed along edges of forest and/or in small forest patches. 4 Migrant Statuses: LD = Long distance migration, typically birds breed during the summer in the U.S. and Canada and they migrate south to spend the winter months in Mexico, Caribbean islands, Central America, and South America; MD = medium-distance, typically birds move south of their breeding range but still within the U.S.; SD = short-distance, typically birds move within their breeding range; YR = year-round resident; IR = irregular/irruptive migrant [Allaboutbirds.org, Cornell Bird Lab] 5 Late-Successional Forest = late successional forests where most of the trees that form the canopy are over 30 ft. tall, including both relatively young forests with trees 15 – 50 years old and mature forests with trees 50+ years or older. This indicates the most likely breeding habitat but is not an indication of likelihood of breeding in small patches. For example, if interior forest specialist confidence is “high” and a “yes” for breeding in mature forest, then this species only breeds successfully in large mature forest patches (> 50 ha). 6 Early-Successional Forest = Composed primarily of shrubs (with some scattering of trees and grassland patches) and/or very young planted pine saplings and pioneer species such as black cherry (Prunus sp.). Trees are generally 0-15 years old and tree height is typically less than 30 ft. 6 For example, if interior forest specialist confidence is “high” and a “yes” for breeding in early-successional forest, then this species only breeds successfully in early-successional forest that is embedded in continuous forest patches (> 50 ha). 7 This column indicates whether the species is considered an interior forest specialist during breeding season. “High”, “Med”, and “Low” refer to the confidence on whether the bird is considered an interior forest specialist. “High” means more that 66% of the reviewed studies indicated that a species was an interior forest specialist, “Med” means between 66% and 33% of studies indicated that a species was an interior forest specialist, and “Low” means that less than 33% of studies indicated that a species was an interior forest specialist. “No” means that the species is not an interior forest specialist because three or greater studies consistently found no area sensitivity; thus, it is likely to breed in small forest fragments. A lack of any of these classifications indicates that fewer than three studies were found for this species in Breeding Review, and thus could not be assigned one way or the other as an interior-forest species. Further, a “—“ in this column represents that the species was not observed in any of the Breeding Review studies. Numbers in parentheses represent the number of studies that determined a species to be an interior forest specialist. Numbers in parentheses represent how many studies noted the species to be an interior-forest specialist out of the total number of studies that observed said species. 8 “Breeding-Migration” represents the transitional period between breeding and migration seasons (either spring transitioning to summer or summer transitioning to fall). Some studies in our review did not clearly define when bird observations were recorded and counted species sightings from these studies in this column. 9 We reviewed four studies that identified Synanthropic birds—species that have adapted to living in urban areas. “High”, “Med”, and “Low” refer to the confidence on whether the bird is considered synanthropic. “High” means more that 66% of at least three reviewed studies indicated that a species was synanthropic, “Med” means between 66% and 33% of studies indicated that a species was synanthropic, and “Low” means that less than 33% of studies indicated that a species was synanthropic. “No” means that the species is not synanthropic because three or more studies found that the species did not display synanthropic behavior and is unlikely to breed in within urban areas. Numbers in parentheses represent the number of studies that determined a species to be a synanthropic out of the total number of studies that observed said species. 10 Marks in these four columns are an indication of using the habitat based on looking across the three reviews. For ease of interpretation, we created four “overall” columns at the end of the bird list. These were “Breeds in Forest Fragment”; “Stopover in Forest Fragment”; “Breeds in Residential Area”; and “Stopover in Residential Area.” For a given species, a checkmark (✓) indicates likelihood of occurrence in each columnar habitat category, an (X) indicates that it does not occur in this habitat category, a (?) indicates that a species may occur in this habitat category, but results were not convincing enough to assign a (✓) or (X), and a blank indicates no information available. First, for species only found in the Breeding Review, if they had three or more studies and were given a High/Med confidence level as an interior-forest specialist, then is it was assigned an (X) in the Breeds in Forest Fragment and Breeds in Residential Area categories; it does not breed in forest fragments in rural and urban areas. Low confidence or a total of two or fewer papers in the Breeding Review, we assigned a (?) to indicate that this species may or may not breed in forest fragments in urban and rural areas. For species found only in the Stopover Review, if a migrant had three or more studies, then we assigned (✓) under the Stopover in Forest Fragment category. If a migrant had fewer than three studies in the Stopover Review, then we assigned (?) under the Stopover in Forest Fragment category. For migrants found in both the Stopover Review and Built Environment Review, if Stopover Review studies combined with two or fewer Built Environment Review studies (stopover, breeding, and breeding-stopover studies), which brought the total to three or greater, then we assigned a (✓) for Stopover in Forest Fragment and a (?) for Stopover in Residential Area categories. The rationale here is that migrants seen in two or fewer Built Environment studies receive a (?) for Stopover in Residential Area because that is not enough studies to confidently assign them as using residential areas as stopover sites For species found in the Breeding Review, Built Environment Review, and Synanthropy Analysis, we compared results to determine what to assign in each habitat category. If a species was a High/Medium confidence interior-forest specialist, but it was observed in three or more studies in the Built Environments Review during the breeding season and was shown to have at least one synanthropic study, we assigned a (✓) for this species as Breeding in Forest Fragment and Breeding in Residential Area. If a migrant, we also assigned a (✓) for Stopover in Forest Fragments and Stopover in Residential Area because if it breeds in residential areas we assumed it would use urban forest fragments and trees in residential areas as stopover sites. For species found in Breeding Review, Built Environment Review, but not in the Synanthropy Analysis, a High/Medium/Low or unassigned confidence level for interior-forest specialist that was a year-round resident and/ or a SD migrant, and it occurred in three or more Built Environment breeding studies, we assigned a (✓) as Breeding in Forest Fragment and Breeding in Residential Area. Because they were not migratory, we were confident that the residential studies reflected breeding individuals. If a High/Medium or unassigned confidence level for interior-forest specialist was a long-distance and/or medium-distance migrant species (even if a portion of the population is considered year-round) and occurred in three or more Built Environment breeding surveys, we were conservative and assumed these individuals were passing through cities. We assigned a (✓) for them under Stopover in Residential Area and Stopover in Forest Fragments. However, if a migrant species was determined to be a Low confidence interior-forest specialist and occurred in three or more Built

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asterisk ( ) indicates for that habitat category, we assigned an (X) or (✓) based on information obtained from other habitat categories. For example, if a species was shown through empirical evidence to have • An interior-forest status, then we could assume that it would not breed in highly fragmented residential areas.

Appendix B. Supplementary data Supplementary data to this article can be found online at https:// doi.org/10.1016/j.landurbplan.2019.01.005.

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*

Environment Review breeding studies, we assigned a (✓) for the Breeds in Residential Area and Breeds in Forest Fragment categories because the combination indicates that they may also breed in residential areas. In turn, because they breed in residential areas then they would also tolerate fragmented forests. For species that occurred only in the Built Environment Review and had three or more total studies (combined stopover, breeding, and breeding-stopover studies), we interpreted the likelihood of this species breeding in residential areas in the following way. If they were observed within the built environment during the breeding season and were shown to have at least one synanthropic study, then we assigned a (✓) for Breeding in Forest Fragment and Breeding in Residential Area. If a migrant, we also assigned a (✓) for Stopover in Forest Fragments and Stopover in Residential Area because if they breed in residential areas they would use forest fragments and residential areas as stopover sites. If species were only observed in the breeding season surveys (not in stopover and breeding-stopover studies), we assigned a (✓) only for year-round residents and SD migrants under Breeds in Forest Fragment and Breeds in Residential categories. All long-distance and medium-distance migrants were considered to be individuals that were passing through and were most likely using the areas as a stopover site. Here, we assigned a (✓) for these species for Stopover in Forest Fragment and Stopover in Residential Area categories. Additionally, for long-distance and medium-distance migrants that had three or more combined stopover, breeding, and breeding-stopover studies under the Built Environment Review, we assigned a (✓) for these species for Stopover in Forest Fragment and Stopover in Residential Area categories. For species that occurred only in the Built Environment Review and had fewer than three total studies, we interpreted the likelihood of this species breeding in residential areas in the following way. For year-round resident species and/or SD migrants, we assigned a (?) in the corresponding Breeds in Forest Fragment and Breeds in Residential categories. All long-distance and medium-distance migrants were assumed to be passing through and may be using residential areas as stopover sites. We assigned a (?) for these species for Stopover in Forest Fragment and Stopover in Residential Area categories.

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