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Our study highlights the value of genetic data from offspring in cases where samples were not available; through analysis of offspring genotypes, we were able to reconstruct genotypes of unknown or unsampled breeders for our analysis Table B in S1 File.

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Combining this new information on kinship and genotypes with long-term field data allowed us to characterize several aspects of urban peregrine breeding behavior and dispersal. We found that Midwest urban peregrines show high fidelity to both mates and nest sites. While these behaviors are characteristics of raptors, they have not been studied extensively in populations living in urban environments.

Higher breeding densities [ 31 ], higher productivity [ 13 ], frequent territorial battles [ 8 , 11 ], and variability in nesting structures [ 8 ] are among many factors that might lead to differences in breeding behaviors in urban populations. Nest site fidelity was even higher than mate fidelity, with only six observed changes in nest sites. Thus, our data shows that mate turnover is higher than territorial turnover, supporting Tordoff and Redig [ 10 ] hypothesis regarding apparent durability of pair bonds. There was no significant difference between males and females in nest site fidelity.

Unsuccessful breeders were more likely to switch nest sites than individuals that bred successfully. Given that structure type and substrate has been shown to be correlated with productivity [ 8 ], moving to a different nearby structure could increase chances for successful reproduction. Indeed, five of the six moves were to nearby nest sites.

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While we did not include structure type or substrate in our analysis, Gahbauer et al. Also, direct human assistant by peregrine monitors in returning grounded fledglings to nests may artificially increase reproductive success and nest site fidelity. Meanwhile, there was no significant correlation between nest site fidelity and breeding density.

This suggests that other aspects of nest sites may be more important in breeding success than interference from other breeders for urban peregrines.

Also, when an established breeder at a productive nest site fails to return it is quickly replaced by another bird. Our finding of very high nest site fidelity is in agreement with reports of other raptor species [ 32 ] with a few noteworthy exceptions. Peregrine falcons in northern Spain switched nest sites on average every three years, and were more likely to change nest sites after raising a large brood [ 33 ]. Rotating among alternate sites within a territory is also common in American kestrels [ 35 ].

Accipiters [ 35 ] and eagles [ 36 ] also move nest sites. It would therefore be interesting to compare nest site fidelity in urban peregrines to that of peregrines nesting in more traditional habitats, but to our knowledge the latter has not been investigated.

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Brood size was a good predictor of EPP in this population. Mougeot [ 31 ] found that EPC frequency in raptors was positively correlated with breeding density. These findings suggested that we might find relatively high rates of EPP in urban peregrines. However, we found only two offspring from one nest that could be considered the product of EPP, yielding a rate of only 1.

Further, there were special circumstances associated with this EPP. The tending male lost his mate in and remated with a female likely already impregnated by another male Table 5. Genetic data confirmed that this male nested from until and sired 26 chicks over this time. Although our paternity analysis included males from neighboring nests, we did not find the sire of the EPP chicks, so it is possible that the male involved in the EPP event was a floater.

The only other reported case of EPP in peregrines also was in an urban nest and there were also special circumstances [ 37 ]. An established breeder who lost its territory after a fight established a new territory, and DNA fingerprinting indicated that this male fathered chicks in both the old nest and the new nest. Our finding of a very low rate of EPP agrees with reports in other falcons nesting in natural habitats.

While Falconids have been shown to be more closely related to Passerines than they are to other raptors [ 38 ], they have lower rates of EPP compared to passerine birds. Low EPP in urban peregrines despite increasing breeding density may be explained by parental care and mechanisms of paternity assurance.

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In long-lived species like peregrines and other Falconids in which bi-parental investment is substantial, females may avoid extra-pair copulations because males reduce their breeding effort when their paternity is in doubt [ 43 ]. Our results support the hypothesis that in places where competition for nests and mates is high, sperm competition will also be high [ 31 ]. Low EPFs may be the result of behaviors aimed to ensure paternity such as displays that enhance pair bonding and a high frequency of within-pair copulations.

Given this level of interaction among successful pairs, extra-pair copulations may be costly to pursue in terms of time and risk of injury or death due to interactions with the territorial male. Our findings on natal dispersal distances are in general agreement with those reported by Dennhardt and Wakamiya [ 9 ] and previous studies on the Midwest population [ 10 ].

We looked at a subset of the data Dennhardt and Wakamiya [ 9 ] used and obtained similar estimates of dispersal distance.

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We found the average dispersal distances of females in both studies were km, about twice that for males km here and km in [ 9 ]. Urban-recovered peregrine populations from the Eastern US [ 46 ] and from Northern Spain [ 47 ] also show female-biased dispersal. While there has been variation among studies in estimated dispersal distances for both sexes, the general trend of females travelling further than males is universal. Some individuals usually males settle at their hack or natal site while others usually females move hundreds of km to different urban areas.

The concentration of nest sites or food resources in urban areas may influence variation in dispersal distances. Many individuals may disperse shorter distances, away from natal sites but in the same urban center. Dispersal distances for those that disperse to new urban centers will depend on the proximity of suitable urban habitats [ 9 ]. Urban peregrine falcons offer an important system for continued study, not only because of their remarkable recovery, but also because their populations will continue to adapt to anthropogenic landscapes.

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Peregrine falcons have become highly visible icons of urban wildlife that promote public education and citizen science. Our study was greatly enhanced by combining molecular markers with field observations. Blood samples of banded chicks are being collected every year, and long term field monitoring has produced an extensive, accessible database. Therefore the opportunity exists to continue our approach and to monitor and study the ecology of urban peregrine falcons.

Table B in S1 File. Detailed contribution from field data and genetic data on the kinship relationships for all Midwestern surveyed nests. We thank Kevin A. Feldheim for helping in the development of microsatellites. We are grateful to the curators, collections managers, and their institutions that kindly provided materials for this study: David Willard, Field Museum of Natural History, Robert M.

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This work was supported by Donald L. This work was completed in partial fulfillment of the requirements for the doctoral degree to ICC from the Graduate College, University of Illinois at Chicago. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. National Center for Biotechnology Information , U. PLoS One. Published online Jul Isabel C. Bates , 3 Mary Hennen , 4 and Mary V. Ashley 1. John M. Mary V.

Lorenzo Zane, Editor. Author information Article notes Copyright and License information Disclaimer. Competing Interests: The authors have declared that no competing interests exist. Received Jan 4; Accepted Jun This is an open access article distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

This article has been cited by other articles in PMC. Abstract Peregrine falcons Falco peregrinus were extirpated from most of the continental United States by widespread use of the pesticide DDT in the s. Introduction Urbanization and rapid expansion of the built environment are generally seen as threats to native species and biodiversity. Table 1 Cities, nest sites, samples, and brood composition of Midwest peregrines analyzed in this study, including both genotyped and non-genotyped individuals.

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Open in a separate window. Ethics Statement No research protocol was needed for our study because samples were collected for general use by peregrine state monitors. Microsatellite screening and genotyping We used a modified enrichment protocol [ 17 ] to isolate microsatellites from two Midwestern peregrine falcons. Kinship, breeding and natal dispersal analyses For evaluating our microsatellite genotypes, deviations from Hardy-Weinberg equilibrium, and the presence of null alleles were tested in CERVUS v.

Table 2 Repeat motifs, primers sequences, sizes, and GenBank Accession numbers for 11 peregrine falcons microsatellite markers.