Southeastern Naturalist C.A. Kelly, S.E. Cameron, and A.C. Boynton 2018 Vol. 17, No. 1 130 2018 SOUTHEASTERN NATURALIST 17(1):130–140 Peregrine Falcon Breeding Performance in North Carolina during the 13-Year Post-delisting Period of 2003–2015 Christine A. Kelly1,*, Susan E. Cameron2, and Allen C. Boynton3 Abstract - We monitored the breeding performance (territory occupancy, nest success, productivity) of a reintroduced hybrid population of Falco peregrinus anatum (Peregrine Falcon) in western North Carolina during the 13-y post-delisting period of 2003–2015. Peregrine Falcons nested at 18 sites (17 cliffs, 1 building), 6 of which were newly discovered sites. Eight to 13 territories were occupied annually. We documented 139 nesting attempts and production of 171 young. Mean nest success (55%) during the period 2003–2015 fell below the 1999–2002 national average, but was more than double earlier (1987–1992) efforts in western North Carolina. Likewise, mean productivity (1.23 young/ year/pair) was more than double earlier efforts in North Carolina, but reflects the disproportionately high contribution of ~5 sites. Throughout the study, subadult birds were members of a pair (13.7% of nesting attempts) and had lower nest success than adult pairs. High variability in nest success and productivity underscore the need for continued monitoring and protection from disturbance and other threats, especially at the most vulnerable and least productive sites. Introduction Breeding populations of Falco peregrinus anatum Bonaparte (American Peregrine Falcon) were absent from North Carolina for more than 2 decades. The last documented nesting attempt was in 1957 at Devil’s Courthouse (Berger et al. 1969, Lee 1999). The historic status and distribution of the Peregrine Falcon in North Carolina is not completely known because no systematic survey of the species was completed prior to the loss of the population. Available records suggest the presence of nesting pairs at 10 sites in the North Carolina mountains before extirpation (Brewster 1886, Ganier 1931, Pearson et al. 1942). Recovery entailed listing the Peregrine Falcon as Endangered under the Endangered Species Conservation Act of 1969, the subsequent banning of DDT, and reintroduction efforts by states (USFWS 2003). The founder population for the recovery effort was comprised of several subspecies (Falco peregrinus anatum, F. p. peali Ridgeway, F. p. peregrinus Tunstall); very few of North Carolina’s hacked captive-bred birds were F. p. anatum. In North Carolina, 92 captive-bred nestling Peregrines were released via hacking or fostering at cliff sites beginning in 1984. Of these, 80 were released from 1984 to 1991 and another 12 were released in 1996 and 1997 in response to a drop in the number of territorial pairs in the 1North Carolina Wildlife Resources Commission, 32 Amber Lane, Asheville, NC 28803. 2US Fish and Wildlife Service, 160 Zillicoa Street, Asheville, NC 28801. 3North Carolina Wildlife Resources Commission, NCSU Centennial Campus, 1751 Varsity Drive, Raleigh, NC. 27606. *Corresponding author - christine.kelly@ncwildlife.org. Manuscript Editor: Douglas McNair Southeastern Naturalist 131 C.A. Kelly, S.E. Cameron, and A.C. Boynton 2018 Vol. 17, No. 1 mid-1990s. In 1987, North Carolina Wildlife Resources Commission (NCWRC) began monitoring the growing breeding population annually while reintroductions continued (Boynton and Currie 1993). In August of 1999, the US Fish and Wildlife Service (USFWS) determined that the Peregrine Falcon had recovered and removed it from the List of Threatened and Endangered Species (Mesta 1999). At the time, North Carolina’s breeding population numbered 9 pairs and was listed as Endangered by the state. Federal delisting triggered a post-delisting monitoring effort of no less than 5 y, as required under Section 4(g)(1) of the Endangered Species Act and implemented by states (USFWS 2003). In response to concerns about the long-term future of Peregrine Falcons, the post-delisting monitoring effort was spread out to monitor the species 5 times at 3-y intervals beginning in 2003 and ending in 2015 (USFWS 2003). However, due to concerns about North Carolina’s small population, the NCWRC elected to monitor nests annually. The purpose of this study was to document the breeding performance (territory occupancy, nest success, productivity) of this newly released population for the 13-y post-delisting period of 2003–2015. Methods Field-site description The study area encompassed cliff habitat in the mountains and foothills of 11 counties in western North Carolina and a building in 1 county of the south-central Piedmont. Cliff habitat ranged from sheer granite cliffs to quartzite and granitic domes. Fourteen of the occupied cliffs are situated on public land (67% national forest, 13% national park, 20% North Carolina state park), and the other 3 are on private holdings. We added an urban site to the study area in 2013, with the discovery of a pair of Peregrine Falcons nesting on the outside of the 40th floor of a building in downtown Charlotte. Seasonal or year-round cliff closures are set on public lands to prevent disturbance from recreational use (e.g., rock climbing, hiking). 2003–2015 surveys The focus of Peregrine Falcon monitoring efforts was initially on the 12 sites where pairs had been present during at least 1 of the 5 y between 1998 and 2002 leading up to and immediately following the de-listing. Gradually, we began checking more cliff sites, including unoccupied historic eyries, potentially suitable cliffs, and rock outcrops classified as medium to high quality, based on expert opinion, as well as other sites brought to our attention. As a result, we incorporated occupied sites into the monitoring effort. NCWRC staff and volunteers monitored sites annually, when possible. In some years, due to staffing and time constraints, we limited monitoring efforts to only those sites with seasonal rockclimbing closures in effect. We began nest-monitoring activities each year in late February or early March to determine territory occupancy, and continued into July and occasionally August. Staff and volunteers used binoculars and 30–60x spotting scopes to watch sites from distant-observation points and often relied on behavioral clues to determine Southeastern Naturalist C.A. Kelly, S.E. Cameron, and A.C. Boynton 2018 Vol. 17, No. 1 132 the stage of nesting (Cade et al. 1996). Observers visited a site approximately every 2–6 weeks to document the nesting stage and, ultimately, the number of young. We assigned the age composition of each pair to 1 of 2 categories: adult (consisting of 2 adults) or subadult–adult (consisting of 1 adult and 1 subadult). We identified subadults by their retained brown juvenal remiges and rectrices, vertical barring on the breast, and replaced gray body contour feathers (Ratcliffe 1980). We identified females by their broad upper body, robust size, brownish-gray and ivory plumage, and pale-yellow legs; and males by their slim, compact shape, bluish-gray and white plumage, and bright yellow legs. Whenever possible, observers also noted intraspecific competition such as an unpaired intruder or replacement of a resident bird, interspecific competition (particularly with Corvus corax L. [Common Raven]), presence of predators near a nest, and disturbances. We considered a territory “occupied” if, during at least 1 of two 4-h site visits, either a pair was observed or a single bird exhibited behavior that provided evidence of reproduction by a pair (USFWS 2003). In some cases, where the minimum number of observation sessions could not be completed, we concluded that a territory was unoccupied based on other evidence such as Common Ravens nesting at the usual Peregrine eyrie, Coragyps atratus (Bechstein) (Black Vulture) and/or Cathartes aura (L.) (Turkey Vulture) activity on or near the eyrie, or absence of fresh whitewash (bird droppings), thus minimizing the probability of overlooking the presence of a pair. The small size of North Carolina’s population rendered moot random sampling of territories. Thus, we summarized territory occupancy, nest success, and productivity for all pairs, including mixed subadult–adult pairs and newly discovered pairs. We considered a nesting attempt successful if young reached 28 d of age (USFWS 2003), regardless of whether fledglings were later confirmed. Rarely, we accepted a count of young that were less than 28 d old when we could not return for a follow-up survey (Johnson and Williams 2014). We defined nest success as the percentage of occupied territories with young and productivity as the number of young per occupied territory. Given our broad definitions, we may have overestimated nest success and productivity. We examined field observations to see if the probable cause(s) of nest failure could be inferred. Data analysis We calculated mean occupancy, nest success, and productivity for each site, each year, and for pooled data. We calculated nest success and productivity for all nesting attempts regardless of outcome and for successful nesting attempts only. In keeping with the USFWS monitoring plan, we report 90% confidence intervals around the mean (USFWS 2003). We applied the chi-square contingency statistic with Yates correction to the nesting data to test for independence of pairs’ age composition and frequency of nesting success versus failure. Data were not normally distributed, so we employed a Kruskal–Wallis non-parametric analysis of variance to examine for any differences in productivity among years. We set statistical significance for both tests at α = 0.05. In keeping with our less stringent Southeastern Naturalist 133 C.A. Kelly, S.E. Cameron, and A.C. Boynton 2018 Vol. 17, No. 1 calculations of 90% rather than 95% CI, we also inspected annual mean nest success by year to search for years when 90% CI values lay outside the pooledmean value for nest success. Results Occupancy Peregrine Falcons nested at 18 sites (17 cliffs, 1 building) during the 13-y postdelisting period (Fig. 1). Our monitoring effort began with 12 sites, and we added 6 sites over the course of the study. Two of the 6 sites were new, whereas the provenance of the other 4 sites was uncertain. The pooled mean-occupancy rate was 82% (Table 1). We found 8–13 occupied territories annually (mean = 10.7), although the proportion of sites occupied dropped sharply after 2010 (Table 2). Among years, occupancy varied from a low of 62% in 2011 to 100% in 2004 and 2007 (Table 2). Among sites, occupancy varied from lows of 14% (Victory Wall) and 50% (Hanging Rock) to a high of 100% (8 eyries; Table 3). Only 5 sites were occupied by a pair every year of the study. Peregrine Falcons abandoned sites because of excessive human disturbance (Devil’s Courthouse) and when a pair of Bubo virginianus Gmelin (Great Horned Owl) nested ~60 m below the falcon eyrie (Shortoff Mountain). Subadult birds were members of a pair in 19 of 139 (13.7%) pooled nesting attempts. The proportion of subadult–adult pairs was highest in 2012, when 3 mixed pairs represented 33% of the total, and was 0% in only 3 years (Table 2). Among the subadults, 79% were females, 16% were males, and sex was not determined for the remaining 5%. Frequent replacement of adult by subadult females occurred at White Rock Cliff (n = 4 years) and Big Lost Cove (n = 3 years). Figure 1. The location of Peregrine Falcon breeding sites in western North Carolina during the 13-year post-delisting period of 2003–2015. Site numbers are cross-referenced with site names in Table 3. Southeastern Naturalist C.A. Kelly, S.E. Cameron, and A.C. Boynton 2018 Vol. 17, No. 1 134 Table 1. Peregrine Falcon mean population indices for western North Carolina post-delisting (2003–2015), with 90% confidence intervals versus national (1999–2002) and North Carolina mid-reintroduction (1987–1992). Geographic region Monitoring period (y) Territory occupancy (%) Nest success (%) Productivity References North Carolina Post-delisting (2003–2015) 82 (77–87) 55 (48–62) 1.23 (1.05–1.41) This study National Post-delisting (1999–2002) 84 68 1.2–1.9A USFWS 2003 North Carolina Mid-reintroduction (1987–1992) 46 25 0.52 Boynton and Currie 1993 AMin–max of means over the 4 years. Table 2. Survey effort, number of pairs, number of pairs with subadults, territory occupancy, brood size, nest success, and productivity of Peregrine Falcons in North Carolina by year during the post-delisting period of 2 003–2015, with 90% confidence intervals. # pairs % nest Mean productivity Sites # with a % territory # pairs with brood size Successful success All sites Successful Year surveyed pairs subadult occupancy 0 1 2 3 4 pairs ± 90% CI # young ± 90% CI sites ± 90% CI 2003 11 10 0 91 4 1 3 2 0 6 60 ± 17 13 1.30 ± 0.73 2.17 ± 0.62 2004 11 11 2 100 5 2 2 1 1 6 55 ± 16 13 1.18 ± 0.77 2.17 ± 0.96 2005 11 10 0 91 0 2 1 4 0 7 70 ± 16 16 1.60 ± 0.78 2.29 ± 0.69 2006 12 11 0 92 4 2 2 3 0 7 64 ± 15 15 1.36 ± 0.70 2.14 ± 0.66 2007 13 13 1 100 5 1 4 0 1 6 46 ± 13 13 1.00 ± 0.64 2.17 ± 0.81 2008 14 13 3 93 6 2 3 1 1 7 54 ± 13 15 1.15 ± 0.66 2.14 ± 0.79 2009 14 12 2 86 9 1 0 2 0 3 25 ± 12 7 0.58 ± 0.60 2.33 ± 1.94 2010 14 13 3 93 5 0 7 3 0 8 62 ± 12 17 1.31 ± 0.55 2.13 ± 0.24 2011 13 8 1 62 5 0 3 0 0 3 38 ± 23 6 0.75 ± 0.69 2.00 ± 0.00 2012 14 9 3 64 3 2 2 2 0 6 67 ± 19 12 1.33 ± 0.76 2.00 ± 0.74 2013 13 10 1 77 5 1 2 1 1 5 50 ± 18 12 1.20 ± 0.86 2.40 ± 1.09 2014 14 9 1 64 1 2 2 4 0 8 89 ± 13 18 2.00 ± 0.69 2.25 ± 0.59 2015 16 10 2 63 3 0 2 2 1 5 50 ± 18 14 1.40 ± 0.91 2.80 ± 0.80 Southeastern Naturalist 135 C.A. Kelly, S.E. Cameron, and A.C. Boynton 2018 Vol. 17, No. 1 Nest success Pooled-mean nest success was 55% (Table 1). Among years, nest success was highest in 2014 (89%) and lowest in 2009 (25%) (Table 2); in both years the 90% CI values for nest success did not overlap with the 13-y mean (Fig. 2). Among sites occupied by breeding pairs for a minimum of 5 y, nest success was highest at Whiteside Mountain (92%) and lowest at Big Lost Cove (18%) (Table 3). Nest success was significantly higher for adult pairs than for subadult–adult pairs (χ2 = 8.12, df = 1, P = 0.004). Productivity We documented 139 nesting attempts during the post-delisting period, 77 of which were successful, producing 171 young; thus, pooled-mean productivity was 1.23 young/pair/y at all sites (Table 1) and 2.22 at successful sites. Mean productivity at all sites was not significantly different among years (H = 10.62, df = 12, P = 0.56). Five sites (Whiteside Mountain, Looking Glass Rock, Panthertail Mountain, Shortoff Mountain, Buzzard’s Roost) produced 63% of all offspring (Table 3). The first 3 of these sites have produced 82% of offspring since reintroduction efforts began in the 1980s (NCWRC, unpubl. data). Causes of nest failure We failed to determine the probable cause(s) of nest failure for 30 of the 62 (49%) failed nest attempts (or for 22% of all nest attempts). Among the 32 remaining failed nests, observers noted several potential causes, including: membership of a subadult in a pair (n = 14; 23% of nest failures), intraspecific competition and replacement (n = 6; 9%), interspecific competition with Common Ravens for nest sites (n = 5, 8%), and other factors (11%; e.g., presence of a predator in the vicinity of the nest, injury of resident bird, human disturbance, documented severe storm event). Figure 2. Annual mean nest success of Peregrine Falcons in western North Carolina from 2003 to 2015. Vertical lines represent 90% confidence intervals on the annual means. Dashed horizontal line represents pooled mean nest success (55%) for the 13-year study. Southeastern Naturalist C.A. Kelly, S.E. Cameron, and A.C. Boynton 2018 Vol. 17, No. 1 136 Table 3. Survey effort and reproductive performance at 18 Peregrine Falcon sites in western North Carolina during the post-delisting period of 2003–2015. # years # years % years with Mean productivity # years occupied occupied subadult # years % nest Total # All years All years Site # Site surveyed by pair by pair in pair successful success young occupied successful 1 Big Lost Cove 13 11 85 3 2 18 2 0.18 1.00 2 Hickory Nut Gorge 13 13 100 1 3 23 7 0.54 2.33 3 Devil's Courthouse 13 9 69 0 5 56 7 0.78 1.40 4 Grandfather Mountain 12 8 67 3 2 25 5 0.63 2.50 5 Hanging Rock 8 4 50 0 1 25 1 0.25 1.00 6 Shortoff Mountain 13 9 69 0 6 67 17 1.89 2.83 7 NC Wall 8 8 100 2 3 38 7 0.88 2.33 8 Looking Glass 13 13 100 0 9 69 25 1.92 2.78 9 Panthertail Mountain 13 13 100 1 10 77 20 1.54 2.00 10 Buzzard's Roost 12 10 83 2 6 60 17 1.70 2.83 11 White Rock Cliff 13 13 100 5 7 54 12 0.92 1.71 12 Whiteside Mountain 13 13 100 2 12 92 28 2.15 2.33 13 Dunn's Roc 7 4 57 0 1 25 2 0.50 2.00 14 Table Rock 3 2 67 0 1 50 3 1.50 3.00 15 Victory Wall 7 1 14 0 1 100 2 2.00 2.00 16 Pickens Nose 5 4 80 0 4 100 7 1.75 1.75 17 Laurel Top 1 1 100 0 1 100 2 2.00 2.00 18 Charlotte 3 3 100 0 3 100 7 2.33 2.33 Southeastern Naturalist 137 C.A. Kelly, S.E. Cameron, and A.C. Boynton 2018 Vol. 17, No. 1 Discussion Over the course of our study in western North Carolina from 2003 to 2015, we discovered new Peregrine Falcon nest sites and documented re-use of some historic sites. Territory occupancy, nest success, and productivity have all substantially improved compared to the mid-reintroduction phase (1987–1992). Territory occupancy was similar to the national mean during 1999–2002 (USFWS 2003). Consequently, we consider occupancy during the post-delisting period in North Carolina to be generally acceptable, although the proportional decline after 2010 warrants further investigation. Our results for nest success were variable and lower than the national mean. Differences in nest success across years evidently occurred in 2009 (low) and 2014 (high). These results coincided with high total rainfall in western North Carolina from March to May 2009 and the rare contribution of young from 2 sites (Hickory Nut Gorge, NC Wall) in 2014. High nest-success in North Carolina was primarily associated with ~5 sites, including 3 sites that were occupied during every year of the study. Regularly occupied territories are successful more often than irregularly occupied territories (Steenhoff and Newton 2007, Wightman and Fuller 2006). Two conspicuous exceptions in our study were the Big Lost Cove and Hickory Nut Gorge sites, which were infrequently successful. Productivity of 1–2 young/pair/y should result in at least a stable population (USFWS 2003). Based on this metric, our mean-pooled productivity rate (1.23) in North Carolina was acceptable, but productivity was boosted by high performance at ~5 sites. Brambilla and Bionda (2013) observed a similar pattern in a Bubo bubo L. (Eurasian Eagle Owl) population in Italy, where 3 of 10 monitored territories contributed disproportionately to productivity, masking problems with nest success at other sites. In California, Peregrine Falcons that bred in rural areas were less productive than those at urban sites (Kauffman et al. 2003). We only have very limited comparable data in North Carolina, where the recently established urban breeding pair at Charlotte was successful all 3 years it was surveyed. Due to our limited vantage points at many eyries and long gaps between site visits, we could infrequently ascertain the exact cause and at what stage a nesting attempt failed. We observed the presence of factors known to lower reproductive performance as well as known sources of disturbance to raptors even if not directly linked to nest failure in our study. Nonetheless, our limited data suggest that the presence of a subadult bird as a member of a resident pair was the most important factor contributing to nest failure. We observed mixed subadult–adult pairs throughout the study, with regular turnover of individuals. Most mixed-age pairs were comprised of a subadult female and an adult male. Yearling females are more frequently a member of a pair than yearling males, possibly a result of yearling males lacking the hunting skills required to feed a mate and young (Wendt and Septon 1991). However, the success of yearling females in a pair is only marginally better than males, and these nesting attempts are characterized by failure to produce eggs or by smaller clutches (Wendt and Septon 1991). Nest failure by subadult females at White Rock Cliff was compounded by competition with nesting Common Ravens in 2014 and 2015. In raptors, a steady number of mixed subadult–adult pairs over Southeastern Naturalist C.A. Kelly, S.E. Cameron, and A.C. Boynton 2018 Vol. 17, No. 1 138 time may suggest a problem with adult survival (Sánchez-Zapata et al. 2000) that may be an early warning sign of ensuing decline (Ferrer et al. 2003). Other than the presence of inexperienced subadult individuals, human disturbance and other factors during the nesting cycle may reduce nest success, such as the simultaneous occurrence of Common Ravens and climbers, which depressed nest success and productivity of Peregrine Falcons in the Alps (Brambilla et al. 2004). Hickory Nut Gorge, NC, was occupied by a pair of Peregrine Falcons during 19 consecutive years, but failed most years. The cliff face is closed to rock climbing, but hiker use increases as the nesting season progresses (C.A. Kelly, pers. observ). In 2015, we observed a helicopter fly near the cliff during the nesting season. Visitors entering a closed area atop Devils Courthouse was a documented disturbance to Peregrine Falcons, and they had abandoned the site by 2012. The NCWRC and National Park Service (NPS) launched a campaign to reduce disturbance. In 2016, NPS staff posted new signs along the trail, conducted outreach via social media throughout the nesting season, and increased law-enforcement patrols and citations for unlawful behavior. Peregrine Falcons subsequently returned and nested successfully. Management recommendations Since efforts to reintroduce breeding Peregrine Falcons began in North Carolina in the 1980s, their recovery has been encouraging. Our study highlights areas for improvement of breeding performance and provides direction for monitoring and management. The subset of sites with regular occupancy by adult breeding pairs, and high nest-success and productivity are our highest conservation priorities. However, lower breeding-performance at many sites remains a concern. Factors contributing to nest failure that can be controlled, such as human disturbance, should be mitigated. The NCWRC has the responsibility in statute to develop a conservation plan for Peregrine Falcons and may petition agencies or private entities to request assistance in implementing conservation, protection, or restoration measures that are beyond the scope of the NCWRC’s authority (NCGA 2010). Currently, the US Forest Service manages seasonal closures during the nesting season, the NPS manages year-round closures, and North Carolina State Parks use a mixed strategy of closures. Eyries on private property are protected at the discretion of the landowner. We recommend that land managers improve closure signage and increase enforcement of closures, but acknowledge that agencies have limited lawenforcement resources to dedicate to such patrols. As demonstrated at Devil’s Courthouse, a surge in education and enforcement can make a positive impact that can be revisited as recreationists fall back into unacceptable habits. Use of helicopters and unmanned aerial vehicles during nesting season should cease. We recommend that land managers establish closed airspace around occupied cliffs during the nesting season. Increased monitoring, including the use of cameras, may be needed to elucidate causes of nest failure at some low-performing sites (e.g., Big Lost Cove). Consistent, comprehensive monitoring coverage of sites is needed to improve accuracy and precision of observations and reveal causes of nest failure. Visiting sites more frequently could be accomplished through a structured volunteer program Southeastern Naturalist 139 C.A. Kelly, S.E. Cameron, and A.C. Boynton 2018 Vol. 17, No. 1 that provides training, lets volunteers adopt cliff-monitoring duties, loans spotting scopes to volunteers, and enlists the aid of rock climbers. At sites with frequent, unexplained nest failure, we recommend posting trail cameras and dataloggers, and checking nests as soon as possible following severe weather events. A high proportion of breeding subadult birds may have implications for the demography of the population due to their lower productivity than adults (Pandolfi et al. 2004). The continued occurrence of inexperienced subadults as members in pairs may indicate problems with adult survival. Band observations or returns from 6 Peregrines hacked in North Carolina during restoration showed return to the natal site and dispersal to the North Carolina Piedmont, US Gulf Coast, and western Mexico (NCWRC, unpubl. data). Research that improves our knowledge of the full life cycle of North Carolina’s Peregrine Falcons is needed. Locational information via band resighting or GPS-transmitters would be useful to determine patterns of mortality outside of the breeding season, especially for yearling and adult females. Acknowledgments This monitoring effort would not have been possible without the volunteers and agency personnel who spent countless hours monitoring nests: L. Appling, D. Bennett, L. Bilbrey, S. Bosworth, L. Brandon, D. Brown, L. Bruce, P. Carroll, J. Catlin, R. Cooke, S. Cooke, I. Evretjarn, A. Frisbee, K. Gemmer, S. Gilland, T. Gordon, K. Gunther, A. Hendrickson, M. Hopey, A. James, J. Johnson, J. Mays, J. Mitchell, R. Mitchell, D. Landwehr, C. McGrath, T. McManus, K. Parker, E. Poole, J. Pope, S. Reed, D. Rupp, Z. Rhodes, N. Schubert, J. Schwierjohann, S. Skeate, A. Steed, G. Taylor, K. Thomas, E. Walker, K. Weeks, M. Westphal, L. Williams, and A. Wilson. Special thanks to J. Haas, North Carolina Outward Bound School, and Southern Appalachian Raptor Research. Finally, we appreciate M. Sará’s and B. Millsap’s helpful reviews of the manuscript. This study was initiated and coordinated by the NCWRC, and funding was provided by State Wildlife Grants and Federal Aid in Wildlife Restoration. The findings and conclusions in this article are those of the authors and do not necessarily represent the views of the USFWS. Literature Cited Berger, D.D., C.R. Sindelar, Jr., and K.E. Gamble. 1969. The status of breeding Peregrines in the eastern United States. Pp. 165–173, In J.J. Hickey (Ed.). Peregrine Falcon Populations: Their Biology and Decline. University of Wisconsin Press, Madison, WI. 596 pp. Boynton, A.C., and R. Currie. 1993. Productivity of reintroduced Peregrine Falcons in western North Carolina. Proceedings of the Annual Conference of the Southeastern Association of Fish and Wildlife Agencies 47:386–393. Brambilla, M., and R. Bionda. 2013. Variation in productivity and territory occupancy in an Eagle Owl, Bubo bubo, population. Ornis Fennica 90:50–56. Brambilla, M., D. Rubolini, and F. Guidali. 2004. Rock climbing and Raven (Corvus corax) occurrence depress breeding success of cliff-nesting Peregrines (Falco peregrinus). Ardeola 51:425–430. Brewster, W. 1886. An ornithological reconnaissance in western North Carolina. Auk 3:94–112. Cade, T.J., J. Enderson, and J. Linthicum (Eds.). 1996. Guide to Management of Peregrine Falcons at the Eyrie. The Peregrine Fund, Boise, ID. 97 pp. Southeastern Naturalist C.A. Kelly, S.E. Cameron, and A.C. Boynton 2018 Vol. 17, No. 1 140 Ferrer, M., V.P. Penteriani, J. Balbontı́n, and M. Pandolfi. 2003. The proportion of immature breeders as a reliable early warning signal of population decline: Evidence from the Spanish Imperial Eagle in Doñana. Biological Conservation 114:463–466. Ganier, A.F. 1931. Nesting of the Duck Hawk in Tennessee. Wilson Bulletin 43:3–8. Johnson, T.H., and S.O. Williams III. 2014. Declining productivity of nesting Peregrine Falcons in New Mexico, 2001–2013. Southwest Natural History Institute, LaJara, NM. 17 pp. Kauffman, M.J., W.F. Frick, and J. Linthicum. 2003. Estimation of habitat-specific demography and population growth for Peregrine Falcons in California. Ecological Applications 13:1802–1816. Lee, D.S. 1999. Extinction, extirpation, and range reduction of breeding birds in North Carolina: What can be learned? Chat 63:103–122. Mesta, R. 1999. Endangered and threatened wildlife and plants; final rule to remove the American Peregrine Falcon from the Federal list of endangered and threatened wildlife. Federal Register 64:46,542–46,558. North Carolina General Assembly (NCGA). 2010. North Carolina Code. Chapter 113 Conservation and Development. Article 25: Endangered and Threatened Wildlife and Wildlife Species of Special Concern. 113–333. Powers and duties of the Commission. Available online at http://www.ncga.state.nc.us/gascripts/statutes/statutelookup. pl?statute=113-333. Accessed September 2017. Pandolfi, M., G. Gaibani, and A. Tanferna. 2004. Depicts the number of breeding pairs reliably the status of Peregrine Falcon (Falco peregrinus) populations? Ardea 92:247–251. Pearson, T.G., C.S. Brimley, and H.H. Brimley. 1942. Birds of North Carolina. Edwards and Broughton Printing Co., Raleigh, NC. 380 pp. Ratcliffe, D.A. 1980. The Peregrine Falcon. Buteo Books, Vermillion, SD. 416 pp. Sánchez-Zapata, J.A., J.F. Calvo, M. Carrete, and J.E. Martínez. 2000. Age and breeding success of a Golden Eagle, Aquila chrysaetos, population in southeastern Spain. Bird Study 47:235–237. Steenhoff, K., and I. Newton. 2007. Assessing nesting success and productivity. Pp. 181–192, In D.M. Bird and K.L. Bildstein (Eds.). Raptor Research and Management Techniques. Hancock House Publishers, Surrey, BC, Canada. 464 pp. US Fish and Wildlife Service (USFWS). 2003. Monitoring plan for the American Peregrine Falcon, a species recovered under the Endangered Species Act. US Fish and Wildlife Service, Division of Endangered Species, Migratory Birds, and State Programs. Pacific Region, Portland, OR. 53 pp. Wendt, A.M., and G.A. Septon. 1991. Notes on a successful nesting by a pair of yearling Peregrine Falcons (Falco peregrinus). Journal of Raptor Research 25:21–22. Wightman, C.S., and M.R. Fuller. 2006. Influence of habitat heterogeneity on distribution, occupancy patterns, and productivity of breeding Peregrine Falcons in central West Greenland. Condor 108:270–281.