JUST, Vol. IV, No. 1, 2016 Trent University Urban Raptors: Owl and Hawk Adaptation to Urban Centers Matthew Poppleton Abstract This report focuses on the adaptation of raptors, specifically hawks and owls, to urban environments. Sighting data was gathered from 3 online sources using Peterborough based data. In the city of Peterborough, the broad-winged, red-tailed, sharp-shinned hawks (Buteo platypterus, Buteo jamaicensis, and Accipiter striatus respectively) and the great gray and great horned owls (Strix nebulosi and Bubo virginianus respectively) are recorded in high numbers relative to other species. The ability for raptors to adapt to urban settings is dependent on ecological limiting factors, including prey items and habitat availability. Many owls and hawks seem to adapt well to urban settings, however there are variances within species and local-populations. Maintaining sufficient natural resources within urban centers is necessary to conserve healthy raptor populations. Keywords Ecology — Ornithology — Environmental Science Introduction The ecology of urban wildlife is a relatively recent research field, which aims to provide wildlife conservationists and managers with knowledge of how animals respond to urban environments. This information can also be used by city planners to build and maintain effective resources and developments to facilitate the continued use of urban centers by animals. In this report, raptors, specifically owls and hawks, are of interest because they are a charismatic species that people have admired for millennia (Kaufman 2000). Urban wildlife management is more effective when focusing on a species that citizens want to protect. Protecting and managing these raptor species also necessitates the protection of wildlife that raptors depend upon, species which otherwise may be thought of as pests, including small mammals, rodents and birds. This article contains a primary report and a literature review. The primary report provides sighting data on owls and hawks within Peterborough, Ontario, and discusses the findings. The literature review discusses the current and past literature on raptor adaptation to urban centers with respect to habitat and diet; urban hazards and recommended management directions are also discussed. The report aims to be relevant to Peterborough, however these implications are relevant for other urban centers as well. 1. Primary Report: Peterborough Hawk and Owl Sightings 1.1 Methods: Raptor Sighting Sources I collected hawk and hawk sighting information from three separate sources: the Cornell Lab of Ornithology and National Audubon Society’s eBird, the Peterborough Field Naturalist’s Christmas Bird Count (CBC), and the Atlas of the Breeding Birds of Ontario (ABBO). This section discusses the different sources and their methodologies. The website eBird.com is a citizen science tool, in which individuals, organizations, and publications with registered online accounts can input bird species sightings from certain locations including Peterborough (eBird.org 2016a). The bird count and date is included with each recorded sighting, along with options of photos and notes. Sighting totals for each species are not available so the individual highest counts of each species from specific dates are used instead. This source of sightings is particularly useful for urban birds because individuals can enter sightings from any location within the city (eBird.org 2016a); the other sources can only record birds from set locations within the city. An issue with eBird is that although historical dated sightings are available, active citizen science records only begin around 2012-2013, creating a disproportionate amount of recent sightings to past sightings (eBird.org 2016b; 2016c; 2016d). Another source of data are The Peterborough Field Naturalist’s (PFN) Christmas Bird Count (CBC), a local chapter of The National Audubon Society’s data collection (Peterboroughnature.org 2015a). The sighting data is recorded by registered groups of volunteers located at separate locations along a 24 km diameter within Peterborough County during one day in December (Peterboroughnature.org 2015b). It should thus be noted that each year records one day’s worth of sightings. Online data is available from the Audubon Society from 1900-2014 (Netapp.audborn.org 2016). I totalled sightings over the past 20 years. Urban Raptors: Owl and Hawk Adaptation to Urban Centers — 2/12 The Atlas of the Breeding Birds of Ontario (ABBO) record uses a point count system in which birds are counted within specific regions. The dataset includes information on breeding evidence and various point count indices, and so the methodology may be more systematic than the CBC and eBird; however, I did not find the actual methodology used. ABBO’s data is more limited by time than the other two sources because it only has data available from 1981-1985 and 2001-2005 (Birdsontario.org 2016). Also, ABBO and the CBC may be more representative of species within specific areas than eBird, because they show trends for birds appearing within set locations, however, they have limited timescales. eBird is not restricted by time, since the birds can be located anywhere within Peterborough at any time, and the boundaries are much larger than the plots of the CBC and ABBO. Since each of the sources used have different timescales and recording methodologies, the counts cannot be totaled to show anything in particular other than the relative number of owl and hawk sightings in Peterborough. 1.2 Sighting Data and Discussion Before synthesizing the results, it needs to be understood that all of the recording methods only demonstrate the number of birds that can be recognized by the recorders. Sightings are likely influenced by recognizable traits, spatial and temporal flight patterns as well as and bird calls produced noise. Due to these issues, the sighting data can only demonstrate the presence and absence of certain species; presence confidence should increase with higher numbers because they are less likely to be mistakes (Table 2; Table 3). The data may also be used as a rough estimate of urban abundance based on the count of each species, however the above issues need to be taken into account (Table 2; Table 3). Additionally, information of the relative amount of raptor urban use and degree of use from Hager (2009) is included. Table 3 compares sighting trends across Peterborough, Toronto and Waterloo. Waterloo is selected due to similarities of population size and area to Peterborough (Table 1); there are no other urban centers in Ontario similarly sized to Peterborough within eBird’s database (Statcan.gc.ca 2016a; 2016b). Toronto is included due to its massive land area and population, and its proximity to Peterborough (Table 1) (Statscan.gc.ca 2016c). Comparing the eBird highest counts from Peterborough with Waterloo and Toronto shows some species trends: the broad-winged hawk (Buteo platypterus) is the bird with the highest hawk count for all three regions, with notably higher counts than the rest of the hawk data (Table 3). However, only eBird in Peterborough shows high results for the broadwinged hawk (Table 2). The consistently high broad-winged hawk counts may be the result of sightings during migratory periods, in which these hawks can be found in flocks with thousands of individuals (Allaboutbirds.org 2016a). Additionally, the red-tailed hawk (Buteo jamaicensis) has the second highest hawk count for all three regions (Table 3), and is the only species (owl or hawk) which has consistently high counts from all three Peterborough sources (Table 2). Redtailed hawks have been described as “the most seen hawks in Ontario” by Bezener (2000) which may explain the consistently high counts (p 97). The sharp-shinned hawk is the third most recorded hawk for Toronto and Peterborough, but neither of these counts are considerably high (Table 3). The red-tailed hawk and sharp-shinned hawk are rated as a potential urban resident with 3/3 points of degree of urban use, whereas the broad-winged hawk is only rated a regular user with 2/3 points (Table 2; Hager 2009). All three of these hawks use urban areas for the winter, migration and breeding (Table 2; Hager 2009). The Cooper’s hawk (Accipiter cooperii) is also a potential urban resident with a 3/3 score of urban use, but does not have high sighting counts (Table 2; Table 3; Hager 2009). Also, the gray hawk’s (Buteo plagiatus) 2 sightings in Peterborough from the CBC are likely misidentifications, because the gray hawk is a tropical species of southern North America and South America (Allaboutbirds.org 2016b). The 1 Swainson’s hawk (Buteo swainsoni) sighting in Toronto may also be a misidentification, because its normal distribution, similar to the gray’s hawk does not extend to Ontario (Allaboutbirds.org 2016c). The owl species sightings are all below a count of 100, and do not follow any location trends (Table 2; Table 3). The absence of barn owl (Tyto alba) sightings from every source and location except Waterloo (count of 1), is likely indicative of its endangered status (Ontario.ca 2016). The two owls with notably higher sightings from the rest of the owls are the great gray owl (Strix nebulosa; 83 sightings from eBird) and the great horned owl (Bubo virginianus; 40 sightings from the CBC) (Table 2). The great gray owl likely has high counts because it is the tallest and has the greatest wingspan of all North American owls and is thus relatively easy to identify (Allaboutbirds.org 2016d). The great horned owl is also a large owl that is also easy to identify, due to the ear-like feathered tufts that on its head (Allboutbirds.org 2016e). The low overall owl sightings may be more due to the difficulties of studying owl habituation due to their quiet nocturnal activity, migration and habitat loss (Dykstra et al. 2012; Hager 2009). 2. Literature Review and Discussion: Urban Raptor Adaptation 2.1 Limiting Factors and Generalists/Specialists It may seem reasonable that the human disturbances, low vegetation levels and structures associated with urban centers would limit the productivity and nesting sites of bird species; however, many published studies have found that many birds, including raptors, adapt well to urban habitats (including: Hager 2009; Dykstra et al. 2012; Chase and Walsh 2006; Rodewald and Kearns 2011; Stout et al. 2006; Minor et al. 1993; Boksakowski and Smith 1997; Campbell 2009). The ability of a species to successfully adapt to urban areas is controlled by certain ecological limiting factors. Animals living in urban areas will typically be affected by different Urban Raptors: Owl and Hawk Adaptation to Urban Centers — 3/12 Table 1. The population and land areas of the City of Peterborough, Waterloo and Toronto, using information from Statistics Canada’s 2011 Census. Statscan.gc.ca 2016a; 2016b; 2016c respectively. 2011 Census Population 2011 Census Land Area (km2 ) Peterborough Waterloo Toronto 78,698 63.8 98,780 64.02 2,615,060 630.21 limiting factors than in natural areas due to altered ecosystems and resource availability (Adams et al. 2006). Adams et al. (2006) suggest that there are generally fewer limiting factors for wildlife within urban centers than natural habitats due to an increased abundance of resources including food, water and shelter, and decreased predator abundance. Raptors are limited by both the availability of habitat for nesting and food/prey resources, and may be generalists or specialists with respect to both these resources (Bird et al. 1996; e.g. Rullman and Marzluff 2014). A specialist species requires certain prey to meet its dietary needs or certain habitat needs, while a generalist is able to use a broader variety of prey/habitats to meet the same needs. Prey availability is influenced by prey vulnerability and predatory access (Rullman and Marzluf 2014). These factors shift across urban environments, and thus shift prey availability (Rullman and Marzluf 2014). Prey vulnerability is affected by many factors including foraging behaviour, bird feeders, breeding status/courtship behaviours, nest selection and overall landscape-based predation access risks (Rullman and Marzluf 2014). Additionally, the density of avian nesting is limited by resource density, quality and by territorial zones from neighbouring birds – which is to say the amount of food, nesting site availability and competition (Newton 1986 as cited in Mannan and Boal 2000). Rullman and Marzluf (2014) observed that prey generalist Cooper’s hawks (Accipiter cooperii) and barred owls (Strix varia) were able to meet dietary needs within urban and forested areas with sufficient land-cover. Thus prey access is likely interlinked with habitat factors. Specialist animal species are usually limited in their ability to adapt to urban centers, due to the specific habitat and/or prey requirements (Gilbert 1989). Campbell (2009) suggests that the potential for specialist species to colonize greenbelts is affected by the degree of similarity between greenbelt forest and contiguous forests, and how recreational activities affect these species. This would likely also apply to specialist species nesting in urban landscapes. Thus, the similarity of urban forests to contiguous forests may affect the degree to which specialist species can adapt to urban habitats. Generalist species are more likely to adapt to urban environments with less strict boundaries. Urban areas contain a relatively low predator abundance, anthropogenic feeding, high perching availability, good access to water bodies along with natural land-cover areas (Campbell 2009) and thus likely attract generalist species. 2.2 Habitat/Nesting The ability of hawks and owls to habituate within urban environments is suggested to be dependent on the populations of species rather than species alone (Dykstra et al. 2012). This is to say that a population of a certain raptor species may be able to thrive within some urban centers and not others. Thus it is important to remember when reviewing the species’ within the literature that the success of a species population to colonize an urban area does not mean that all populations of that species will colonize urban areas (Dykstra et al. 2012). However, reports concerning local population adaption may be used to provide a synthesis of avian responses to urban environments, and more comprehensive ideas can apply to broader regions. 2.3 Land Cover A key factor affecting of bird habitat is the land-cover type (the landscape type on a regional scale: grassland, forested land, developed land, etc.). The number of bird species associated with certain habitats has been found to be higher within Ottawa’s urban centers than greenbelts and some contiguous forestland, due to greater land-cover heterogeneity within the urban areas (Campbell 2009). This is also the case with raptors. Stout et al. (2006) found that red-tailed hawks (Buteo jamaicensis) had greater nesting use of heterogeneous habitats, which are representative of suburban areas, than homogenous habitats representative of highly developed areas. The heterogeneous nesting habitat had over three times the amount of unused grassland and woodland land-cover for nesting (Stout et al. 2006). This may imply that the redtailed hawk may successfully nest within Peterborough due to the city’s large amount of suburban areas, surrounding green spaces and inner parks. These areas may provide sufficient grassland and forested land-cover for red-tailed hawks and other raptor species. Undeveloped land-cover within urban centers can support rare urban birds (Loss et al. 2009). Thus, undeveloped land-cover will likely support habitat-specialist raptors. The availability of natural land-cover patches can also influence the potential of migratory birds using urban habitats (Konze 2009). For information on managing urban raptor habitats see Literature Review: Management Implications. Raptors seem to thrive within high natural and urban land-cover types, as demonstrated in western Washington (Figure 3) (Rullman and Marzluf 2014). Thus the ability for raptors to excel within Urban Raptors: Owl and Hawk Adaptation to Urban Centers — 4/12 Table 2. Hawk sightings in Peterborough Ontario from three separate bird sighting counts. Records are from eBird’s highest individual count records; the Peterborough Field Naturalist’s Christmas Bird Count’s (CBC) totals over the past 20 years; the Atlas of Breeding Birds of Ontario’s (ABBO) records from 2001-2005. Degree of opportunity is on a scale of 1-3, representing inconsistent use, regular use and potential residency, respectively (Hager 2009). Urban activity use describes whether the bird uses urban areas for breeding (B), the winter (W) and for migration (M) (Hager 2009). N/A is included for species that are included by Hager (2009), but are not reported to use urban centers. Degree of urban activity and urban activity use are modified from (Table 1 as cited in Hager 2009, p 214) (Sightings are from eBird.og 2016b; Netapp.auborn.org 2016; Birdsontario.org 2016) Species eBird CBC ABBO Degree of Ur- Urban ban Activity Activity Use Hawk Species Broad-winged hawk (Buteo platypterus) Cooper’s Hawk (Accipiter cooperii) Ferruginous hawk (Buteo regalis) Gray Hawk (Buteo plagiatus) Northern Goshawk (Accipiter gentilis) Red-Shouldered Hawk (Buteo lineatus) Red-tailed hawk (Buteo jamaicensis) Rough-legged hawk (Buteo lagopus) Sharp-shinned Hawk (Accipiter striatus) Swainson’s Hawk (Buteo swainsoni) 362 3 0 2 0 3 10 0 7 0 0 69 0 0 11 0 660 7 51 0 18 1 40 0 3 11 17 0 5 0 3 3 2 1 1 3 3 N/A 2 2 Owl Species eBird CBC ABBO Degree of Ur- Urban ban Activity Activity Use Barn Owl (Tyto alba) Barred Owl (Strix varia) Boreal Owl (Aegolius funereus) Eastern Screech-Owl (Megascops asio) Great Gray Owl (Strix nebulosa) Great Horned Owl (Bubo virginianus) Long-eared Owl (Asio otus) Northern Hawk Owl (Surnia ulula) Short-eared Owl (Asio flammeus) Snowy Owl (Bubo scandiacus) Northern Saw-whet Owl (Aegolius acadicus) 0 13 1 2 83 5 1 1 1 1 11 0 0 0 3 0 40 0 0 1 1 0 0 3 0 0 0 2 1 0 0 0 0 2 2 N/A 3 N/A 2 1 1 1 3 2 certain land-covers seems to be dependent on the species, population, and land-cover type. 2.4 Species and Population Traits Separate species sizes can also associate with separate landcover types. For example, in a study by Campbell (2009), large birds were more commonly found within contiguous forests than small birds. This may be due to high tree density and low human activity (Campbell 2009). Also, generalist species were not found in the contiguous forest areas in this study. This may imply that larger species are generally habitat specialists. Campbell (2009) suggests that bird presences can be well predicted using habitat types, while species presence patterns are better explained by species type. Different communities have varying success with urban habitats and nesting. Minor et al. (1993) found that great-horned owls and red-tailed hawks were able to successfully nest and repro- BWM BW W B WM BW BWM N/A BWM B B BW N/A BW N/A BW W BW M W BWM duce equally well in urban and natural areas. Similarly, the barred owl’s nesting habitat usage has been observed to be unaffected by suburban developments (Dykstra et al. 2012). This instance may be due to the owls having low interactions with humans because of their low detectability and nocturnal activity as mentioned in Primary Report: Sighting Data and Discussion (Dykstra et al. 2012; Hager 2009). Coleman et al. (2002) found no relationships between the quality of habitats and raptor productivity, implying that cover type may be a stronger parameter than habitat quality. Additionally, for an urban area to be a bird nesting source, it is important to have high reproductive success, nest site re-occupancy, and population recruitment, as observed in Milwaukee, Wisconsin, by Stout et al. (2007). Some species have greater selectivity for certain nesting land-cover types. The sharp-shinned hawk (Accipiter striatus) has been observed to nest within coniferous trees in habitats Urban Raptors: Owl and Hawk Adaptation to Urban Centers — 5/12 Table 3. The highest recorded counts of hawk and owl populations from eBird in Peterborough, Toronto, and Waterloo, Ontario (eBird.org 2016b; 2016c; 2016d respectively). Hawk Species Toronto Waterloo Peterborough Broad-winged hawk (Buteo platypterus) Cooper’s Hawk (Accipiter cooperii) Ferruginous Hawk (Buteo regalis) Gray Hawk (Buteo plagiatus) Northern Goshawk (Accipiter gentilis) Red-Shouldered Hawk (Buteo lineatus) Red-tailed hawk (Buteo jamaicensis) Rough-legged hawk (Buteo lagopus) Sharp-shinned Hawk (Accipiter striatus) Swainson’s Hawk (Buteo swainsoni) 7060 18 0 0 0 40 750 1 646 1 1200 9 0 0 1 8 110 30 15 0 362 3 0 2 0 3 10 0 7 0 Owl Species Toronto Waterloo Peterborough Barn Owl (Tyto alba) Barred Owl (Strix varia) Boreal Owl (Aegolius funereus) Eastern Screech-Owl (Megascops asio) Great Gray Owl (Strix nebulosa) Great Horned Owl (Bubo virginianus) Long-eared Owl (Asio otus) Northern Hawk Owl (Surnia ulula) Northern Saw-whet Owl (Aegolius acadicus) Short-eared Owl (Asio flammeus) Snowy Owl (Bubo scandiacus) 0 2 2 6 2 6 10 1 4 1 2 1 37 1 38 10 1 2 0 13 1 2 83 5 1 1 1 15 5 5 17 1 1 of low coniferous composition (Coleman et al. 2002). Also, the red-tailed hawk has been observed to nest specifically in well-covered and mature stands, however, this may be due to predation threats in other land-cover areas rather than being due to preferential habitats (Coleman et al. 2002). Former nesting sites also influence raptor habitat, for some raptors will use the nests of separate species. The barred owl was observed by Dykstra et al. (2012) to frequently nest in recently used nests of red-shouldered hawks (Buteo lineatus) within suburban areas (Dykstra et al. 2012). Time may also be an influencing factor, for the successful nesting sites of Cooper’s hawks was also observed to have rates of reoccupy of periods of over 2 generations (Stout et al. 2007). Additionally, hawks have been suggested to behave with greater efficiency in new nesting areas, with peak improvements occurring within the first 1-2 years (Mannan and Boal 2000). 2.5 Fragmentation Birds living within urban environments have been observed to favour fragmented habitats. Great horned owls (Bubo virginianus) nesting in Connecticut, northern New Jersey, and southeastern New York were observed to select complex fragmented habitats rather than communities in rural habitats (Smith et al. 1999). Great horned owls were even attracted to rural fragmented land-covers due to human developments (Smith et al. 1999). Some raptors are known to excel within fragmented landscapes, including forest edges and human developed landscapes (Rullman and Marzluff 2014). This attraction towards urbanized fragmented landscapes has been suggested to be due to fragmented forests having greater nesting choices and forest openings than contiguous forests, and greater prey items supplemented by urbanization (Smith et al. 1999). Similarly, sharp-shinned hawks have been reported to nest within close proximity to forest openings associated with human activity within and surrounding the city of Montreal (Coleman et al. 2002). 2.6 Timescales The age of neighbourhoods within certain landscapes may also influence nesting. For example, old suburban neighbourhoods (> 75 years) with dense populations have been observed to include old trees that are likely to include cavities of sufficient size for nesting of species such as the barred owl (Dykstra et al. 2012). Also, old neighbourhoods in Chicago, Illinois, have been observed to have high degrees of undeveloped landcover (Loss et al. 2009) which may provide specialist habitat. However, population shifts have also been observed in aging neighbourhoods, including bird species richness decreasing with neighbourhood age, along with community composition changes from originally native and migratory birds to exotic and non-migratory birds (Loss et al. 2009). Thus older neighbourhoods in Peterborough may support greater diversities of Urban Raptors: Owl and Hawk Adaptation to Urban Centers — 6/12 Figure 1. The richness of raptors within a gradient of urban and natural land-cover types within western, Washington, United states. The gray boxes represent species presence. The species are listed from highest site frequency. (Fig. 2. as cited in Rullman and Marzluf 2014, p 7) owl and specialist species, however, species habitation may also depend on the size of the neighbourhood. 2.7 Human-Built Structures Human-built structures also effect raptor populations. Generally, developed fences, hedges and vegetative edges have been suggested to improve the hunting ability of raptors for prey items (Rullman and Marzluf 2014). For management purposes, these strategies may be implemented in cases where local raptors are detrimentally limited by prey. On a smaller scale, it has been suggested raptor species have greater reproductive rates within human-made artificial nests than natural nests (Chace and Walsh 2006). 2.8 Migratory Sites The City of Toronto released a “Literature Review and Data Assessment” of migratory birds in 2009, that likely has implications for migratory birds in Peterborough. In a broad sense, birds may use nesting areas that have been routinely used for generations, as suggested by this review (Konze 2009). Additionally, many temperate and neotropical birds migrate past Toronto on their migration routes. The proximity of Peterborough to Toronto may indicate that Peterborough is also within the migratory range of these species. Toronto was also suggested to be a nesting stop for migratory birds that cross Lake Ontario; similarly Peterborough may be a reliable stop for birds traveling along the Otonabee River (Konze 2009). 2.9 Prey/Diet 2.10 Prey Abundance The ability of a raptor to find prey is of great importance. Chace and Walsh (2006) state that urban areas can positively influence the abundance of raptor prey, which then positively influences raptor abundance (Chace and Walsh 2006). Redtailed and Swainson’s hawks (Buteo swainsoni) are examples of hawks that likely consume these high abundances of urban prey (Chace and Walsh 2006). Additionally, small prey are specifically abundant in urban centers, and thus the raptors that consume small prey are not likely to be limited by diet in urban centres (Chace and Walsh 2006). Similarly, urban Florida burrowing owls (Athene cunicularia floridana) have been observed with significantly different diets from rural Florida burrowing owls, including with high insect diets and more variable prey items (Mrykalo et al. 2009). However, the study only recorded from one site in both urban and rural areas, so this information may not be due to habitat differences (Mrykalo et al. 2009). Examples of raptors which consume small prey are the burrowing owl and screech owl (Megascops). The ferruginous hawk (Buteo regalis) and rough-legged hawk (Buteo lagopus) are examples of raptors that consume larger prey (Chace and Walsh 2006). Therefore, information on rodent size and species found within Peterborough may be useful to further the understanding of the potential of each bird to adapt to developed landscapes. 2.11 Rodent Abundance Hindmarch and Elliot (2015) observed rat prevalence as positively related to urban-land area at barred owl nesting sites within southwestern British Columbia. Barred owl rat predation was so great within urban environments that rats were rated as the most consumed prey item (Hindmarch and Elliot 2015). However, the consumption of rodents in urban areas can put raptors at an increased risk of rodenticide exposure (see Literature Review: Urban Hazards: Disease and Poisoning) (Hindmarch and Elliot 2015). Raptor size can also limit preying ability within in certain developed areas. Scavenging raptors in the Rio Negro and Neuquén provinces of Argentina are known to use roads for scavenging, while the large hunting raptors do not appear to use the roads. This is suggested to be due to the relatively slow takeoff of large raptors, which can increase the threat of roadside collisions and hunting along roads (Lambertucci et al. 2009). In addition to this abundance of prey, urban centers typically have an abundance of anthropogenic waste and supplementary feeding stations which habituated animals can use as food resources (Adams et al. 2006). Food resources in urban centers are often distributed in concentrated locations, which results in the distribution of animals concentrating in these areas. This Urban Raptors: Owl and Hawk Adaptation to Urban Centers — 7/12 clumped distribution pattern is similar to natural environments (Adams et al. 2006). 2007). This may be a major urban predation threat because of the commonly high populations of raccoons. 2.12 Indications of High Prey Abundance In southeastern Arizona, Estes and Mannan (2003) observed the prey delivery rate of Cooper’s hawks to urban nests to be about twice as much as in rural nests. A high rate of prey delivery to nests may indicate an abundance of prey within the respective habitat (Estes and Mannan 2003). An abundance of readily available prey items can also be indicated by hawks that continually deliver prey items to nestlings, while able to vary their hunting effort to meet nestling diet requirements (Estes and Mannan 2003). When there is a high abundance of prey in a given site, hawk hunting locations will likely become dependent on the location of their nests (Mannan and Boal 2000). 3. Urban Hazards 2.13 Predation on raptors Raptor species that prey upon other raptors will likely affect species abundance and distribution. For example, the Cooper’s hawk is a known predator of the sharp-shinned hawk (Coleman et al. 2002). Thus, urban areas with a high abundance of raptor-predating species such as the Cooper’s hawk may cause reductions in the numbers of the preyed-upon raptors such as the sharp-shinned hawk (Coleman et al. 2002). This is not specifically relevant to the available sighting data for these two species within Peterborough, as the sharp-shinned hawk has higher sighting counts than the Cooper’s hawk from 2 out of 3 of the sighting sources within Table 2 and higher sighting counts within every city in Table 3. Nest depredation is also relevant, for raptors within urban centers have been observed as more likely to prey on avian nests in rural areas than urban areas (Rodewald and Kerns 2011). Similarly, Rodewald et al. (2011) observed separations of interrelated connections between breeding birds and nest predators living in urban areas. This may be due to the abundance of additional prey and food resources for predators within urban environments. This separation may increase successful nesting potential in urban environments, for nestlings likely have a higher survival potential with a lack of predation. Supplementary food from bird feeders in particular can attract avian wildlife to specific locations. Some raptors such as the Cooper’s and sharp-shinned hawk may prey upon the birds that are attracted to feeders, and thus are also attracted to these feeders (Allaboutbirds.org 2016f). However, this is not always the case: Roth et al. (2008) found no significant difference in predation from Cooper’s hawks at bird feeders. Feeders can also be detrimental to the birds that use them and raptors (see Literature Review: Urban Hazards: Feeding Stations). Nests within urban areas may also encounter predation from other urban adapted animals such as raccoons. Raccoons may be an undocumented predator to raptor species and their eggs, as they use similar habitats to Cooper’s hawk nesting sites and prey upon similarly sized eggs and birds (Stout et al. 3.1 Collisions While urban centers seem to be beneficial in terms of prey and habitat, they also affect mortality rates and can be hazardous to resident raptor species. Potential hazards include vehicular and window collisions, built structures, artificial lighting, roads and diseases. In a comprehensive review of 86 articles, Hager (2009) discusses urban hazards to raptor populations. The main hazards were window and vehicle collisions and electrocutions; the collisions and electrocutions are substantial hazards in both urban and natural environments. Lighting from human developments can also cause mortalities. In 1996, the Fatal Light Awareness Program (FLAP) stated that Toronto had an annual mortality rate of 732 deaths due to artificial lighting (Ogden 1996). Light towers are also a major cause of bird mortalities, 6.8 million tower-related bird kills annually are estimated within United States and Canada (Longcore et al. 2012). Thus lit antenna towers and other structures within Peterborough may cause mortalities. In addition, rural land changes such as land-expansion can put pressure on wildlife by limiting or removing habitats (Neave et al. 2000). Developing forested land-cover within urban centers would have the same negative effects on nesting raptors; positive changes would likely occur with park restoration, or forest planting. 3.2 Roads Raptors use roads and their surrounding structures for foraging road kills, hovering and hunting prey (Hager 2009). Roads can also disturb bird populations through noise levels, visual disturbance and prey concealment (Bautista et al. 2004). However, in terms of vehicular collisions, Hager (2009) found that moralities were not significantly different between urban and natural environments. Road disturbance to raptor species’ may occur on weekly cycles depending on traffic variations, because scavenging raptor road-occurrence has been observed to decrease during weekends with high traffic loads (Bautista et al. 2004). This may have implications for Peterborough during summer weekends, when tourism is likely highest. 3.3 Feeding Stations Bird feeding stations can cause issues with urban-living bird species. Although these stations provide a concentrated and consistent source of food, the feeders can directly and indirectly cause mortalities. Feeders located near windows can attract birds and can cause window collision mortalities (Klem 1990). Raptors are also susceptible to window collisions for they may prey on birds located at these feeders (Boal and Mannan 1999). Hunters have also been found to use feeding stations to attract raptor species; this may occur at rates higher than currently recorded if the hunting is illegal (Boal and Mannan 1999). Also, feeders have the potential of exposing Urban Raptors: Owl and Hawk Adaptation to Urban Centers — 8/12 birds to Trichomoniasis (Davidson and Nettles 1988 as cited in Boal Mannan 1999) which may transfer to raptors that feed on these birds (see Literature Review: Urban Hazards: Disease and Poisoning below) (Friend and Franson 1999; Boal et al. 1998). 3.4 Disease and Poisoning Urban adapted raptors may be required to cope with various diseases, including Trichomoniasis. Trichomonas gallinae (Trichomoniasis) is an endemic parasite of which doves (columbidae) are the foremost host (Stabler 1954 as cited in Estes and Mannan 2003; Lawson et al. 2011). This can be passed on to raptor species that prey upon small birds such as doves. Ultimately, the spread of Trichomoniasis depends on the degree to which the hosts are present in a raptor’s diet (Boal and Mannan 1999). Another danger is the infusion of seeds with organophosphates as a poison control of feral pigeons; this similarly puts raptors that consume these pigeons at risk of organophosphate poisoning (Boal and Mannan 1999). In addition, as noted above (Literature Review: Prey/Diet: Rodent Abundance), there can be high abundances of small prey such as rats within urban environments (Chase and Walsh 2006; Hindmarch and Elliot 2015). This abundance would increase the risk of raptors becoming exposed to rodenticides (such as anticoagulant rodenticides [AR]) through secondary exposure; rats were the main pathway for AR to barred owls exposure from a study in southwestern British Columbia (Hindmarch and Elliot 2015). 3.5 Human Communities Raptors within urban and suburban environments can be noticed and appreciated by neighborhood communities (Boal and Mannan 1999). However, there have been instances of individuals being intentionally hit by Cooper’s hawks in Tucson, Arizona, due to nest protection and defensive behaviours (Boal and Mannan 1999). Aside from concerns of injuries, the potential threats of these nests within human communities can cause humans to mistrust raptor nests (Boal and Mannan 1999). Thus, educating the communities with an ecological understanding of this behaviour should positively influence local opinions (Boal and Mannan 1999). 4. Management Implications 4.1 Land-cover To effectively manage for each species, the biology of the species, its population and limiting requirements should be understood. In terms of land-cover, it is recommended by Campbell (2009) to plant a variety of forest densities for human tolerant and non-tolerant bird species. Similarly, Coleman et al. (2002) recommends that forest stands within urban centers should be considered raptor habitats and thus be conserved. Additionally, fragmented forest patches within an urban landscape benefit generalist raptors, and thus should be conserved (Rullman and Marzluf 2014). For species with low human tolerance and/or specialist requirements, a sufficient density of undeveloped patches can be estimated with an understanding of each species’ average flight and alert distances (Campbell 2009). Additionally, fenced corridors between undeveloped patches may increase the potential of colonization of species with low human tolerances (Campbell 2009). Shrub patch size and proximity to roads are also important land features to consider when planning because of their relevant consequences of human activity (Campbell 2009). Loss et al. (2009) suggest that natural land-cover patches can increase avian richness within populations over 2 km away on average, and within urban landscapes composed of less than 20% natural land-cover patches (including park and undeveloped land) (Loss et al. 2009). There are 102 parks with varying sizes and levels of forest cover distributed throughout Peterborough, which are generally within 1-3 km of each other, and thus may increase suitable raptor habitat (Figure 4). 4.2 Species size Urban forests and parks can be managed to provide habitat for larger birds that are not normally associated with urban areas. These birds will likely require large areas of contiguous forests (Campbell 2009). Large raptors sighted in Peterborough include the great horned owl, the great gray owl (Strix nebulosa) and the red-tailed hawk. Larger birds may require isolation within these areas, this can be achieved with high density forests and by fencing off certain areas (Campbell 2009). 4.3 Diseases In managing for bird feed related diseases such as Trichomoniasis, Mannan and Boal (1999) recommend the regular cleaning of feeders, “dove-proof feeders”, and the general avoidance of attracting doves to the feeders (p 82). There are available treatments for Trichomoniasis, however, they usually result in bacterial infections or mortalities even when successful, and thus euthanizing infected individuals has been recommended (Boal and Mannan 1999). A potential alternative of rodenticide use is to use raptors as a natural pest control (Hungeryowl.org 2016). This is also a potential alternative to the use of organophosphate poisons. For example, owls may be encouraged to act as natural rodent pest management by building owl boxes for nesting (Hungryowl.org 2016). Community groups and organizations within Peterborough could advertise and create incentives to implement these within Peterborough. However, in order for this to be effective, poison use would first need to be reduced to prevent exposure, and raptor habitats may require proper management of land-planning. 4.4 Planning Management should focus on specific specialist species, for generalist species are less likely to require management due to their ability to adapt to a greater range of habitats. Within generalists and specialists, it seems that individual species have Urban Raptors: Owl and Hawk Adaptation to Urban Centers — 9/12 Figure 2. Park locations within the City of Peterborough, Ontario, Canada. (For an enlarged map, see Peterborough.ca 2016) Urban Raptors: Owl and Hawk Adaptation to Urban Centers — 10/12 their own requirements for habitat and prey. Therefore, urban managers focused on raptors should manage for specialist species that are affected by the most limiting factors. Additionally, planners may focus on increasing the available habitat for endangered species such as the barn owl, however, this would expose them to potentially threatening urban hazards (see Literature Review: Urban Hazards) (Ontario.ca 2016). Managing for specialist species may create an umbrella of protection, by providing a greater amount of available resources for less limited specialist species and generalist species. In other words, managing for specialists would likely improve resource availability for low abundance raptor populations. However, there is insufficient sighting information for Peterborough to determine which species are of high or low urban abundance. Therefore a systematic evaluation of urban avian wildlife abundance and distribution within the city would be useful for this purpose. 5. Conclusion The ability of raptor species to adapt to urban environments seems to be dependent on resource availability, commonly in terms of habitat and diet requirements and their generalist or specialist requirements with respect to these resources. Generalist species are commonly better able to adapt to urban environments, while some specialist species can adapt with the availability of specific habitat, and prey. Urban center resources can differ from natural areas through factors such as supplementary feeding and prey abundance (Smith et al. 1999; Adams et al. 2006; Hindmarch and Elliot 2015), builtstructure influences (Hager 2009; Rullman and Marzluff 2014; Longcore et al. 2012), decreased contiguous forest cover, natural predators (Campbell 2009; Adams et al. 1006) and increased land-cover heterogeneity (Campel 2009). The Peterborough sighting counts (Table 2 and 3) are limited in their ability to accurately demonstrate distribution or abundance; however, they are important tools for understanding the presence of specific species. The broad-winged and red-tailed hawks had the highest counts of all recorded species, while the great gray and great horned owls had slightly higher counts than other owls. Additionally, there was only one sighting of the endangered barn owl (Ontario.ca 2016). It would be useful to have an intensive record of bird species abundance and distribution in addition to these sightings for Peterborough and other urban centers. With respect to the reviewed literature, many studies have been done in the United States on raptors in urban centers, however, few have been conducting regarding Canadian urban centers. Also, many of the studies focus on how specific species responded to urban habitat/prey availability. This is generally useful information, however, the degree to which these species will show similar trends between separate populations and other urban centers is uncertain. This uncertainty arises because the degree to which a species’ response to urban factors may ultimately depend on the traits of a specific species population rather than the traits of a species in general (Dykstra et al. 2012). This report demonstrates general responses of raptors to urban areas, and it would be useful for other studies to include broader implications on urban responses to seek out more generalized trends. Local results are useful in respect to the local management of species but may limit the usability of their results on a broader scale. A generalized understanding of raptor response to urban areas is also useful, for it provides resources to regions distant from the original study locations. 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