Variability in spatial behaviour and migratory strategies selection in a global warming context: the case of the Hen Harrier at the southern limit of its distribution

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This study investigates the non-breeding or posreproductive movements of Hen Harriers using GPS/GSM data from 30 individuals over 54 periods from 2019 to 2023. Using maximum and mean distances from the reproductive centroid as behavioural proxy, we classified posreproductive periods into in three strategies: migration (22.2%), sedentarism (9.3%), and sedentarism with posreproductive movements (68.5%). Our results challenge the previous belief that the majority of the Spanish Hen Harrier population was sedentary. The study establishes a relationship between distance travelled and latitude of breeding areas, suggesting environmental adaptation. The Spanish Hen Harrier population, being at the southern limit of its breeding range, exhibits less pronounced differences between sexes or strategies. Raptors with extensive migratory and post-breeding periods are prone to be more vulnerable due to the shortening of the area available for posreproductive movements. Animal Science Behavioral Ecology Animal Behavior Animal behaviour Breeding Dispersive migration GPS/GSM Migration Raptor Sedentarism Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction In order to better understand animal ethology, we tend to classify behavioural patterns into different categories or strategies in the different parts of the annual cycle. In the case of birds and in the posreproductive period, they are usually classified into sedentary (or resident) species and migratory species or populations. However, these two classic categories often mask much greater behavioural variability, including intraspecific ones. Newton ( 2010 ) classified movements, accounting for into six different groups, taking into account distance travelled, time and direction: routine day-to-day movements, dispersal movements, dispersive migration, migration, irruptions, and nomadism. The Hen Harrier ( Circus cyaneus ) is a medium-sized raptor with a wide distribution in the Paleartic. This species is originating from the Asian steppes, which has invaded Europe and the Iberian Peninsula over the past three thousand years. The deforestation of wooded areas has provided an ideal habitat for this raptor and other steppe species, which are now much more abundant. Currently, in northern Europe Hen Harriers breed in semi-natural upland areas, even in forest plantation gaps (Amar and Redpath, 2005 ; Amar et al., 2008 ). While in the southern edge of its distribution, in Spain, they breed in pseudo-steppe landscapes, predominantly in cereal crops (Arroyo, 2022 ). Species inhabiting the margins of the distribution tend to show greater variability in strategies and spatial behaviour (Brown, 1984 ; Brown and Lomolino, 1998 ). The Iberian Peninsula, due to its geographical location, is an area that for some species is at the southern limit of its distribution and it is therefore very interesting to study the behaviour of the populations that inhabit it. This occurs in species such as Red Kite ( Milvus milvus ) (García-Macía et al., 2021 ; 2022a ; 2022b ; 2022c ; 2022d ) or Hen Harrier, showing a peculiar behaviour in Iberia, as previous studies suggested (García and Arroyo, 2001 ; Redpath et al., 2002 ; Arroyo, 2022 ; Morollón et al., 2024 ). It is frequent to find Iberian species that are phylogenetically closely related but perform different movement strategies, one of them displaying sedentary tendencies and the other one fully migratory: Hen Harrier and Montagu’s Harrier ( Circus pygargus ) or Red Kite and Black Kite ( Milvus migrans ) (Cramps and Simmons, 1979 ; Ferguson-Lees and Christie, 2001 ; Arroyo et al. 2019 ; García-Macía et al., 2021 ). The closest species to the Hen Harrier, the Montagu's Harrier which also inhabit the Palaearctic and is strictly a long-distance migrant, travelling from their breeding grounds in northern Europe and Asia to the southern hemisphere to spend the winter (Limiñana et al., 2012 ; Panuccio et al., 2014 ). In this study, we present the case of the Hen Harrier in the Iberian Peninsula, where its migratory behaviour displays particular characteristics associated with its position at the southern limit of its distribution in a global warming context. This population includes sedentary, migratory, and dispersive individuals, allowing observation of a range of behaviours that may be shaped by natural selection processes. This case offers a unique opportunity to understand how environmental pressures (such as global warming), phenotypic plasticity, and evolutionary selection interact in the evolution of migratory behaviour. In this study GPS-tracked 30 Hen Harrier to analyse the different posreproductive behaviours of the species. We aimed to 1) define and analyse post-breeding movements using the distances travelled during post-breeding periods; 2) classify any observed differences in posreproductive periods into different movement strategies; 3) study the year-on-year variability of spatial strategies in specific individuals; 4) Provide a tool and database to see the impact of global warming on the post-reproductive strategies of a species in the southern part of its distribution. Materials and Methods Tagging and tracking The movements of 30 adult or subadult Hen Harrier individuals (19 females and 14 males) were monitored with GPS/GSM transmitters from 2019 to 2023. The individuals were captured mainly from central and northern Spain in different provinces: Alava (6), Asturias (2), Cantabria (1), La Rioja (1), Lerida (1), Madrid (2), Salamanca (5), Segovia (3), Toledo (6), Valladolid (3) (Figure S1 and Table S1 in Supplementary Material). These 30 individuals provided us data for 54 posreproductive periods (Table S1 in Supp. Mat.). The individuals were, at least, two years old (del Hoyo et al., 1994) in their first period considered (Euring codes 5 and 6). All individuals reproduced successfully in Spain. The breeding season in the Hen Harrier lasts from March to July (García-Macía et al., unpublised report ) so we analysed the movements in the posreproductive periods, which were generally from August to February, although the dates of the individual reproductive periods were accounted for (Table S1 in Supp. Mat.). Where exact dates were not known, the mean breeding dates of this species were considered (from March 21 st to July 4 th ; García-Macía et al., unpublised report ). Between 2019 and 2022, the Hen Harriers were lured with a Eurasian Eagle-Owl and captured with dho-gaza nets. During the marking process, individuals were ringed and morphometric measurements were taken. Sexing was based on the clear sexual colour dimorphism of the species (Ferguson-Lees and Christie, 2001). Individuals were fitted with satellite tracking transmitters (9g Milsar GSMTag-U9 or 10g-12g Ornitela OrniTrack 10/E10 3G GPS/GSM), back-mounted using thoracic Teflon harnesses. Following established guidelines, the weight of the backpacks was less than 3% of the body weight of each individual (Kenward, 2001). The handling time until release of captured individuals was less than one hour to avoid stressing the animals. The devices were set to collect locations every 15–30 min, depending on battery voltage, season, and time of day. Data packages were transmitted via the digital cellular network. Then the data were filtered with a 30 min period to standardize the sample. Capture of individuals was carried out by expert licenced ringers from the Spanish Ornithological Society (SEO/BirdLife), complying with ethical standards in the capture and tagging of Hen Harriers. The location data of all individuals were provided by SEO/Birdlife and Natural England to the Vertebrate Zoology Research Group of the University of Alicante. Strategies classification Before conducting the analyses, we undertook a qualitative classification of the posreproductive periods of the individuals (Figures 1 and 2). This classification was based on previous studies and reviews (García-Macía et al., 2022c; Newton, 2010), and these strategies can be described as: Migration (M): Individuals exhibiting two distinct latitudinal movements (in spring and autumn) during the posreproductive period between breeding and wintering areas. Where the latitudinal distance exceeds 400 km (Kerlinger 1989: Morollón et al., 2024). Sedentarism : Individuals that did not perform two clear latitudinal movements in the posreproductive period and remain in the breeding area. Sedentarism sensu stricto (S): Individuals that remained very close to the nest for the entire year, without significant displacements. They only performed routine day-to-day movements throughout the year. Sedentarism with posreproductive movements (SM; or dispersive migration): Individuals which performed all-direction movements far away from their nests after breeding season, and then a return movement toward the starting point in time for the next breeding season. It involved both wandering movements and settling in temporary settlement areas (TSAs). These movements have a component more exploratory and dispersive, as is shown in previous works (Rosier et al., 2006; Moss and Hipkiss, 2014; Wheat et al., 2017; Heggøy et al., 2021), similarly to the dispersal movements of immature birds (<2 years old). Spatial parameters Based on previous studies on posreproductive movements (McCaslin et al., 2020; García-Macía et al., 2022c) two variables were considered to analyse the movements of each period: maximum distance from the reproductive centroid (km; MaxD), that is, the Euclidean distance of the farthest location from the reproductive centroid and the mean Euclidean distance from the reproductive centroid (km; MeanD). The centroid of the breeding area was calculated using the smallest weekly 50% kernel of each individual each year. This smallest kernel indicates the time at which the member of the pair were incubating eggs, as we can see in other species of raptors (López-López et al., 2021; Morollón et al., 2022; García-Macía et al., 2023) and therefore the locations are focused on the nest of each pair. To calculate these variables, we used the following R packages: adehabitatHR (Calenge, 2006) to obtain the smallest breeding area, the package rgeos (Bivand and Rundel, 2021) to calculate the breeding centroid and the base R package stats (R Core Team, 2021) to compute the distances during the posreproductive period from the breeding centroid. Statistical analysis We used a framework of Linear Mixed Models (LMMs) to check the differences between these three strategies (M, S and SM) and sexes. In both models, Sex and Strategy were fixed effects, while ID and Period were considered as random effects. The response variables were the Maximum distance to the breeding centroid and the Mean distance to the breeding centroid , with both variables square root transformed to improve model fit. In order to evaluate the significance of mixed effects models, ANOVA tests with Kenward-Roger approximations were performed. Finally, pairwise comparisons using Estimated Marginal Means test (EMM) with Kenward-Roger approximations and with Tukey test method to p-value adjustment were used to find if there are some differences between strategies. Then, we performed a Fisher’s exact test (non-parametric test) to analyse the sex distribution among the three different strategies. Finally, we computed a linear regression between the MaxD and the breeding centroid latitude to know if there are any correlation between the position in the Iberian Peninsula of the breeding area and the movements of the Hen Harriers. Statistical analyses were performed with R software version 4.2.1 (R Core Team, 2021), and the level of significance was set at P<0.05. We used the following R packages: LMMs were computed using lme4 (Bates et al., 2015), ANOVA was calculated with car (Fox and Weisberg, 2019), and the pairwise comparison EMM test with emmeans (Lenth et al., 2022). Linear regression and Pearson’s coefficient were calculated with the package ggpubr (Kassambara, 2020). QGIS software version 3.22.6 (QGIS Development Team, 2022) was used to visualize the breeding areas, centroids and distances of each individual, as well as for the creation of the maps. Results Strategies classification The 54 posreproductive periods (36 from 18 females and 18 from 12 males) were classified into three strategies: Migration (M), Sedentarism (S) and Sedentarism with posreproductive movements (SM) (Figure 3). The most common strategy was SM, with 37 periods (68.5%), 26 females and 5 males; followed by M with 12 periods (22.2%), 8 females and 4 males; and finally, S with only 5 periods (9.3%), 2 females and 3 males (Table 1). Spatial parameters The S strategy had the lower values in both variables, MaxD and MeanD. On the contrary, M had the higher values, and in between these two limits we found the SM (Table 1). We found significant differences in ANOVA test in both variables when we compared the different strategies (MeanD: F=12.22, p<0.001; MaxD: F=13.90, p<0.001; Table 2). In addition, the EMM post-hoc analysis showed these differences in all comparisons except when comparing SDM and SM (Table 3). When we compared both variables between sexes, we observed that there were no differences in MeanD (F=0.498, p=0.488) neither in MaxD (F<0.001, p=0.976) (Table 2). Also were no differences when we compared both Sex and Strategy together MeanD (F=0.15, p=0.870) neither in MaxD (F=0.047, p=0.954) There were no differences in the distribution between Sexes in the different Strategies in the Fisher’s exact test (p>0.05 in all strategies; Figure 4). The linear regressions in the different strategies between the MaxD and the breeding centroid latitude were not significative (all p>0.05), however there were trends in the strategies; migration strategy (M) showed a positive Pearson’s coefficient between both variables (R=0.413), while the sedentary strategies S and SM showed a negative relationship (R=-0.022, R=-0.263, respectively) (Figure 5). Discussion For the first time, the posreproductive strategies of the Spanish population of the Hen Harrier have been studied and defined. The analyses found behaviour that exhibits three different posreproductive strategies: migration with a latitudinal movement of more than 400 km; sedentarism sensu stricto ; and sedentarism with posreproductive movements or also called dispersive migration (Newton, 2010 ). These patterns are the same to those found in the Red Kite in previous studies (García-Macía et al., 2022c ). The most common strategy in Red Kites was sedentarism with 75% of posreproductive periods while migrants accounted for only 10% (García-Macía et al., 2022c ). In Hen Harriers, the most common strategy was sedentary with posreproductive movements with 68.5% and the least common was sedentary with only 9.3% of posreproductive periods. This result is novel as it was previously believed that the vast majority of the Hen Harrier population in Spain did not perform large movements and was sedentary (del Hoyo et al., 1994 ; García and Arroyo, 2001 ; Redpath et al., 2002 ; Arroyo, 2022 ). One explanation for these patterns could be that in addition to the Common Vole ( Microtus arvalis ), Hen Harriers during the breeding season often feed on chicks and fledglings of other birds before they reach full flight capacity (del Hoyo et al., 1994 ; Redalph et al., 2001; Arroyo and Redpath, 2009 ; Ludwig et al., 2018 ). The young of many open-nesting species have a prolonged period of following their parents without reaching full fledging ability, unlike forest species. The lack of this easy resource after the breeding season may lead to a different hunting behaviour, which motivates the search for new food sources, preferably rodents, which is a more dispersed and difficult resource, and consequently a greater dispersal. The results show variability in the behaviour of individuals, which may be a logical adaptation to different environmental, trophic and habitat conditions. For example, harriers fly through areas of recently harvested or unharvested cereal or scrubland and this implies different prey that may also vary, such as population explosions of Common Voles or young fledglings. This adaptive plasticity occurs even in the same individuals that change their strategy from one year to the next (As occurs with individual 200452, which remains sedentary during the first two years and in the third year undertakes a migration of almost 950 kilometres to Morocco; see Table S1 in Supp. Mat.), as is also the case of the Red Kite in the Iberian Peninsula (García-Macía et al., 2022c ). Our results also showed that migration strategies have a connection between distance travelled and latitude of the breeding area, as is the case with partial (in lower latitudes) and full migrations (in higher latitudes) throughout Europe. Most of the breeding areas of Hen Harriers that show shorter movements are located further south than those with larger movements. This may be due to the fact that the birds that breed in the Iberian Peninsula do not have as many ranges of movement compared to those that breed in northern Europe or America, which are characterised by longer migrations. From an evolutionary perspective, the behaviour of the Hen Harrier in the Iberian Peninsula can be interpreted as the result of selective pressures specific to the edge of its geographical range. These pressures tend to favour flexibility in movement patterns (Brown, 1984 ; Brown and Lomolino, 1998 ), enabling individuals to optimise their balance between site fidelity for breeding, resource acquisition during winter, and the reduction of migratory costs. Full migration could be costly in terms of energy and risks associated with movement, whereas complete sedentarism could increase local competition or the risk of food shortages. The behavioural flexibility observed in the Hen Harrier in Spain might represent conditions similar to the origins of migratory behaviour (Salewski and Bruderer, 2007 ). The increasingly benign climatic conditions of the Iberian Peninsula due to the global warming, its position at the southern limit of the species' range, and other factors, enable the coexistence of sedentary, migratory, and dispersive individuals. This variability provides a 'natural laboratory' where selection can act, favouring behaviours most adaptive to the species as selective pressures shift (Berthold, 1999 ; Winkler, 2005 ; Salewski and Bruderer, 2007 ). The most widely accepted hypothesis about the origin of migration suggests that this behaviour evolved gradually from local or dispersive movements, which over time increased in distance and complexity (Salewski and Bruderer, 2007 ). In this context, the characteristics observed in the Hen Harrier may reflect early stages in the evolution of migration or, alternatively, a partial reversion towards residency in response to favourable environmental conditions. In both cases, natural selection seems to be shaping a diversity of movement strategies, ranging from full migration to more flexible and complex behaviours. Site fidelity for breeding is a common trait among all individuals in this population, suggesting that this characteristic may have a strong genetic basis or be favoured for its role in reproductive success (Pulido et al., 1996 ; Pulido and Berthold, 2003 ). However, the variability in migratory behaviour raises questions about the degree of genetic inheritance of these strategies. Although we lack data on kinship relationships to assess whether sedentary individuals originate from sedentary parents or if migrators have migratory progenitors, studies on other species, such as the Red Kite, have shown that these behaviours can change even within the same generation in response to environmental conditions (García-Macía et al., 2022c ), as observed in the Hen Harrier with ID 200452. This case highlights how selection can act rapidly to adjust the behaviour of a population to local conditions. If climatic conditions remain favourable, migratory behaviours may gradually be replaced by sedentary strategies, which are considered more energy-efficient. However, intermediate or hybrid behaviours, such as dispersive migration observed in the Hen Harrier, could also be selected between these extremes. In conclusion, this example provides insight into the natural selection processes shaping the evolution of migratory behaviour in birds. The coexistence of multiple strategies within a single population underscores the importance of plasticity and variability as key factors in adapting to dynamic environments. As environmental conditions continue to change, these selective processes will persist in defining the optimal behaviour for the population. We propose that such conditions are those required for the emergence or evolutionary selection of migratory or sedentary behaviours within a population, as may have occurred in other species during other periods in range-limit areas. Declarations Animal Ethics Statement This study did not involve experimental procedures on animals and therefore, according to Spanish legislation, did not require approval from an animal ethics committee. The research involved only the capture and tagging of wild Hen Harriers (Circus cyaneus) with lightweight GPS devices, which is regulated by regional wildlife authorities in Spain. All trapping and marking activities were conducted under permits issued by the competent regional administrations: • Departamento de Industria, Transición Energética y Sostenibilidad, Gobierno Vasco • Consejería de Administración Autonómica, Medio Ambiente y Cambio Climático, Principado de Asturias • Consejería de Fomento, Ordenación del Territorio y Medio Ambiente, Gobierno de Cantabria • Consejería de Agricultura, Ganadería y Medio Ambiente, Gobierno de La Rioja • Departamento de Desarrollo Rural y Medio Ambiente, Gobierno de Navarra (Permit No. 0001-0261-2020-000027) • Departamento de Acción Climática, Alimentación y Agenda Rural, Generalitat de Cataluña • Consejería de Medio Ambiente, Licencias de Caza y Pesca, Comunidad de Madrid • Consejería de Fomento y Medio Ambiente, Junta de Castilla y León (Permit No. AUES/CYL/28/2021) • Consejería de Agricultura, Ganadería y Desarrollo Rural, Gobierno de Castilla-La Mancha Captures were carried out by expert licensed ringers from the Spanish Ornithological Society (SEO/BirdLife), following strict ethical standards for the capture and tagging of wild raptors. Acknowledgements The results provided in this publication were obtained thanks to the work of SEO/BirdLife in the Migra program and its funding by Fundación Iberdrola España and Naturand England. Numerous individuals, ornithological groups, ringing groups, research centers, forestry agents and technicians from the autonomous communities were involved. Without their collaboration it would not have been possible to carry out part of the work. Many people from SEO/BirdLife also took part in the work, especially Javier de la Puente, Ana Bermejo and Carmen Fernández. And of course the staff who helped with other administrative and logistical elements of project management. From Natural England we would especially like to thank for their great collaboration in the capture of individuals to Javier de la Puente, Arturo Rodríguez and Juan Martínez. Funding This work was supported by Fundación Iberdrola España (MIGRA program of SEO/BirdLife) and Natural England. Data Availability Statement All data used in this study are publicly available upon request to data managers in the online data repository Movebank (www.movebank.org). The projects are: “Hen Harrier in Spain – Migra Program in Spain” (project ID: 1169461367) and “Hen Harriers in Spain” (project ID: 1135251191). Authors’ contribution S.M., and V.U. conceived the ideas and designed the methodology, S.L. managed the data. S.M. and J.G.M. analysed the data. S.M. and V.U. wrote the manuscript. S.M., JGM., S.L. and V.U and contributed critically to the drafts. S.L. and V.U. gave final approval for publication. Conflict of interest The authors declare that no conflict of interest exists. References Amar, A. and Redpath, S.M. (2005). Habitat use by Hen Harriers Circus cyaneus on Orkney: implications of land-use change for this declining population. Ibis. 147 : 37-47. Amar, A., Arroyo, B., Meek, E., Redpath, S. and Riley, H. (2008). 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In: Berthold P, Gwinner E, Sonnenschein E (eds) Avian migration. Springer, Berlin Heidelberg New York, pp 53–77 Pulido F, Berthold P, van Noordwijk AJ (1996) Frequency of migrants and migratory activity are genetically correlated in a bird population: evolutionary implications. Proc Nat Acad Sci USA 93:14642–14647 QGIS Development Team. (2022). QGIS Geographic Information System. Open Source Geospatial Foundation Project. http://www.qgis.org R Core Team (2021). R: A language and environment for statistical computing. RFoundation for Statistical Computing, Vienna, Austria. URL https://www.R-project.org/. Redpath, S.M., Arroyo, B.E., Etheridge, B., Lekie, F.M., Bowman, K. and Thirgood, S. (2002). Temperature and Hen Harrier productivity: from local mechanisms to geographical patterns. Ecography. 25 : 533-540. Redpath, S.M., Clarke, R., Madders, M. and Thirgood, S.J. (2001). Assessing raptor diet: comparing pellets, prey remains, and observational data at Hen Harrier nests. Condor. 103 : 184-188. Rosier, J.R., Ronan, N.A., Rosenberg, D.K. (2006). Post-breeding Dispersal of Burrowing Owls in an Extensive California Grassland. The American midland naturalist. 155 (1): 162-167. Salewski, V., & Bruderer, B. (2007). The evolution of bird migration—a synthesis. Naturwissenschaften , 94 , 268-279. Wheat, R.E., Lewis, S.B., Wang, Y., Levi, T. and Wilmers, C.C. (2017). To migrate, stay put, or wander? Varied movement strategies in bald eagles ( Haliaeetus leucocephalus ). Movement Ecoogy. 5 : 9. Winkler, D.W. (2005) How do migration and dispersal interact. In: Greenberg R, Marra PP (eds) Birds of two worlds . Johns Hopkins University Press, Baltimore, 401-413. Tables Table 1: Percentages, Mean, Standard Deviation and Range of Maximum and Mean distances to the breeding centroid by strategies. Strategy Sex No. of posreproductive periods (%) Mean distance to the centroid (km) Maximum distance to the centroid (km) Migration Total 12 (22.2%) 318.24 ± 231.74 (44.1 - 795.52) 679.27 ± 368.7 (120.55 - 1227.91) Females 8 (14.8%) 292.83 ± 241.47 (44.1 - 795.52) 673.22 ± 352.69 (120.55 - 1082.34) Males 4 (7.4%) 369.06 ± 236 (155.14 - 623.22) 691.38 ± 455.98 (209.28 - 1227.91) Sedentarism sensu stricto Total 5 (9.3%) 8.89 ± 6.02 (3.34 - 19.04) 31.94 ± 24.18 (15.59 - 74.68) Females 2 (3.7%) 8.2 ± 0.36 (7.94 - 8.45) 21.89 ± 1.61 (20.75 - 23.03) Males 3 (5.6%) 9.36 ± 8.47 (3.34 - 19.04) 38.64 ± 31.62 (15.59 - 74.68) Sedentarism with posreproductive movements or dispersive migration Total 37 (68.5%) 127.54 ± 78.86 (7.27 - 448.21) 254.58 ± 153.66 (74.95 - 893.34) Females 26 (48.2 %) 114.36 ± 52.15 (20.01 - 235.14) 250.46 ± 117.5 (101.05 - 560.26) Males 11 (20.3%) 158.71 ± 118.61 (7.27 - 448.21) 264.33 ± 224.36 (74.95 - 893.34) Table 2: ANOVA test results based on the previous LMMs. df: degrees of freedom, df.res: degrees of freedom of the residuals. Significative differences are shown in bold. Variable Factor F df gl.res p-value Mean Distance to Breeding Centroid (Intercept) 85.76 1 27.09 <0.001 Sex 0.56 1 24.41 0.460 Strategy 12.22 2 34.66 <0.001 Sex: Strategy 0.15 2 35.64 0.870 Maximum Distance to Breeding Centroid (Intercepto) 141.60 1 22.86 <0.001 Sex 0.001 1 19.61 0.974 Strategy 13.90 2 26.94 <0.001 Sex: Strategy 0.047 2 27.79 0.954 Table 3: Pairwise comparisons with EMM test of the four strategies. LDM: Large-distance migration, SDM: Short-distance migration, S: Sedentarism, SM: Sedentarism with posreproductive movements. SE: Standard Error, df: degrees of freedom. Significative differences are shown in bold. Variable Contrats Estimador DE df t-ratio p-value Mean Distance to Breeding Centroid M-S 14.89 2.5 45.1 6.06 <0.001 M-SM 6.96 2 25.1 3.56 0.004 S-SM -7.93 2.2 46.1 -3.69 0.002 Maximum Distance to Breeding Centroid M-S 19.93 3.2 36.5 6.33 <0.001 M-SM 9.61 2.1 20.7 4.63 <0.001 S - SM -10.32 2.8 36.9 -3.71 0.002 Additional Declarations The authors declare no competing interests. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6445538","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":445761816,"identity":"302c79f7-24b0-4dcc-8069-ee61ebfd544c","order_by":0,"name":"Sara Morollón","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA2UlEQVRIiWNgGAWjYLCCBwVQxgcgZmMnRkuCAYRmnAHSwkyKFmYeMElAtW5778MHCQY2efLtZx9+tvm1TZ6PmYHxw8cc3FrMzhw3NkgwSCs2OJNuLJ3bd9uwjZmBWXLmNjxabqSxSSQYHE7cwJDGIJ3bc5sRqIWNmZcYLfP7nzH/tuy5bU+8lgYgQ5rhx+1EwlrOHGMG+SVxw41nbJa9DbeT25gZm/H75Xgb44MPFTZAh6Ux3/jx57bt/Pbmgx8+4tGCChjbwGQDsepB4A8pikfBKBgFo2CkAADFeU7TLzL0zQAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0003-4156-3398","institution":"Universidad de Alicante","correspondingAuthor":true,"prefix":"","firstName":"Sara","middleName":"","lastName":"Morollón","suffix":""},{"id":445761817,"identity":"c476263c-2bdb-4353-9c2b-81ad32bfcc81","order_by":1,"name":"Jorge García-Macía","email":"","orcid":"https://orcid.org/0000-0001-5634-5153","institution":"Universidad de Alicante","correspondingAuthor":false,"prefix":"","firstName":"Jorge","middleName":"","lastName":"García-Macía","suffix":""},{"id":445761818,"identity":"a035fd24-8880-451e-8303-53ea290054ff","order_by":2,"name":"Simon Lee","email":"","orcid":"","institution":"Natural England","correspondingAuthor":false,"prefix":"","firstName":"Simon","middleName":"","lastName":"Lee","suffix":""},{"id":445761819,"identity":"25d4e8fa-b19c-46ca-816d-9ffe41e3d6c2","order_by":3,"name":"Vicente Urios","email":"","orcid":"https://orcid.org/0000-0001-6444-379X","institution":"Universidad de Alicante","correspondingAuthor":false,"prefix":"","firstName":"Vicente","middleName":"","lastName":"Urios","suffix":""}],"badges":[],"createdAt":"2025-04-14 11:26:27","currentVersionCode":1,"declarations":{"humanSubjects":false,"vertebrateSubjects":true,"conflictsOfInterestStatement":false,"humanSubjectEthicalGuidelines":false,"humanSubjectConsent":false,"humanSubjectClinicalTrial":false,"humanSubjectCaseReport":false,"vertebrateSubjectEthicalGuidelines":true},"doi":"10.21203/rs.3.rs-6445538/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6445538/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":81937517,"identity":"600c1e19-ad2d-497c-955a-83dc2368b619","added_by":"auto","created_at":"2025-05-05 06:28:35","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":40916,"visible":true,"origin":"","legend":"\u003cp\u003eQualitative classification criteria to delimit the three posreproductive strategies.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-6445538/v1/494a060b792803e9f2a1ab5d.png"},{"id":81937513,"identity":"af3be24d-433c-401a-af46-8deaa6783406","added_by":"auto","created_at":"2025-05-05 06:28:35","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":41045,"visible":true,"origin":"","legend":"\u003cp\u003eScheme of strategies followed by Hen Harriers. A) Migration: Two annual and latitudinal movements of more than 400 km between breeding areas and wintering quarters. B) Sedentarism \u003cem\u003esensu stricto\u003c/em\u003e: day-to-day movements home-range bounded in the whole annual cycle. C) Sedentarism with posreproductive movements or dispersive migration: a combination of wandering movements and stopping in temporary settlement areas (TSA) after breeding season, without marked seasonality or direction, involving a return journey after that in time to next breeding season.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-6445538/v1/8d8283fbddff9474f3089dd6.png"},{"id":81937518,"identity":"f0561023-78da-46c4-9db9-4510289be87e","added_by":"auto","created_at":"2025-05-05 06:28:35","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1987454,"visible":true,"origin":"","legend":"\u003cp\u003eTracking periods of posreproductive strategies.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eA) The 12 posreproductive periods of 8 Hen Harrier doing Migration (M) back and forth to their wintering areas. The example of the latitudinal graph belongs to ID 190217.\u003c/p\u003e\n\u003cp\u003eB) The 5 posreproductive periods of 4 Hen Harrier doing Sedentarism (S) staying in the same place. The example of the latitudinal graph belongs to ID 200428.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;C) The 37 posreproductive periods of 18 Hen Harrier doing Sedentarism with posreproductive movements (SM) with nomadic movements without a specific direction sometimes staying in temporary settlement areas. The example of the latitudinal graph belongs to ID 200424.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-6445538/v1/924d76923924a587935b650b.png"},{"id":81937514,"identity":"cc1f77ad-b693-4943-bfab-54914a99eba7","added_by":"auto","created_at":"2025-05-05 06:28:35","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":65523,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution and percentages of periods between sexes in the different posreproductive strategies. P-values of Fisher’s exact test. M: Migration, S: Sedentarism, SM: Sedentarism with posreproductive movements or dispersive migration.\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-6445538/v1/2931ada8c69ad7ff80a766fa.png"},{"id":81938357,"identity":"b50171ad-5837-4b1e-8537-9f00292180b0","added_by":"auto","created_at":"2025-05-05 06:36:35","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":232051,"visible":true,"origin":"","legend":"\u003cp\u003eLinear regression between the Maximum distance (km) of each individual in each posreproductive period and the Latitude of the centroid (m). M: Migration, S: Sedentarism, SM: Sedentarism with posreproductive movements or dispersive migration, R: Pearson’s coefficient, p: p-value.\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-6445538/v1/0dab3e7bb67399695b999e3b.png"},{"id":81939953,"identity":"b352d0a1-9edb-412a-bc3f-6abe30de9276","added_by":"auto","created_at":"2025-05-05 06:52:41","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3263277,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6445538/v1/b67072c8-3d19-4939-bce7-072885072eef.pdf"}],"financialInterests":"The authors declare no competing interests.","formattedTitle":"Variability in spatial behaviour and migratory strategies selection in a global warming context: the case of the Hen Harrier at the southern limit of its distribution","fulltext":[{"header":"Introduction","content":"\u003cp\u003eIn order to better understand animal ethology, we tend to classify behavioural patterns into different categories or strategies in the different parts of the annual cycle. In the case of birds and in the posreproductive period, they are usually classified into sedentary (or resident) species and migratory species or populations. However, these two classic categories often mask much greater behavioural variability, including intraspecific ones. Newton (\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) classified movements, accounting for into six different groups, taking into account distance travelled, time and direction: routine day-to-day movements, dispersal movements, dispersive migration, migration, irruptions, and nomadism.\u003c/p\u003e \u003cp\u003eThe Hen Harrier (\u003cem\u003eCircus cyaneus\u003c/em\u003e) is a medium-sized raptor with a wide distribution in the Paleartic. This species is originating from the Asian steppes, which has invaded Europe and the Iberian Peninsula over the past three thousand years. The deforestation of wooded areas has provided an ideal habitat for this raptor and other steppe species, which are now much more abundant. Currently, in northern Europe Hen Harriers breed in semi-natural upland areas, even in forest plantation gaps (Amar and Redpath, \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Amar et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). While in the southern edge of its distribution, in Spain, they breed in pseudo-steppe landscapes, predominantly in cereal crops (Arroyo, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSpecies inhabiting the margins of the distribution tend to show greater variability in strategies and spatial behaviour (Brown, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Brown and Lomolino, \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). The Iberian Peninsula, due to its geographical location, is an area that for some species is at the southern limit of its distribution and it is therefore very interesting to study the behaviour of the populations that inhabit it. This occurs in species such as Red Kite (\u003cem\u003eMilvus milvus\u003c/em\u003e) (Garc\u0026iacute;a-Mac\u0026iacute;a et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e; \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e; \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2022c\u003c/span\u003e; \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2022d\u003c/span\u003e) or Hen Harrier, showing a peculiar behaviour in Iberia, as previous studies suggested (Garc\u0026iacute;a and Arroyo, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Redpath et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Arroyo, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Moroll\u0026oacute;n et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIt is frequent to find Iberian species that are phylogenetically closely related but perform different movement strategies, one of them displaying sedentary tendencies and the other one fully migratory: Hen Harrier and Montagu\u0026rsquo;s Harrier (\u003cem\u003eCircus pygargus\u003c/em\u003e) or Red Kite and Black Kite (\u003cem\u003eMilvus migrans\u003c/em\u003e) (Cramps and Simmons, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1979\u003c/span\u003e; Ferguson-Lees and Christie, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Arroyo et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Garc\u0026iacute;a-Mac\u0026iacute;a et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The closest species to the Hen Harrier, the Montagu's Harrier which also inhabit the Palaearctic and is strictly a long-distance migrant, travelling from their breeding grounds in northern Europe and Asia to the southern hemisphere to spend the winter (Limi\u0026ntilde;ana et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Panuccio et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn this study, we present the case of the Hen Harrier in the Iberian Peninsula, where its migratory behaviour displays particular characteristics associated with its position at the southern limit of its distribution in a global warming context. This population includes sedentary, migratory, and dispersive individuals, allowing observation of a range of behaviours that may be shaped by natural selection processes. This case offers a unique opportunity to understand how environmental pressures (such as global warming), phenotypic plasticity, and evolutionary selection interact in the evolution of migratory behaviour. In this study GPS-tracked 30 Hen Harrier to analyse the different posreproductive behaviours of the species. We aimed to 1) define and analyse post-breeding movements using the distances travelled during post-breeding periods; 2) classify any observed differences in posreproductive periods into different movement strategies; 3) study the year-on-year variability of spatial strategies in specific individuals; 4) Provide a tool and database to see the impact of global warming on the post-reproductive strategies of a species in the southern part of its distribution.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003e\u003cstrong\u003e\u003cem\u003eTagging and tracking\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe movements of 30 adult or subadult Hen Harrier individuals (19 females and 14 males) were monitored with GPS/GSM transmitters from 2019 to 2023. The individuals were captured mainly from central and northern Spain in different provinces: Alava (6), Asturias (2), Cantabria (1), La Rioja (1), Lerida (1), Madrid (2), Salamanca (5), Segovia (3), Toledo (6), Valladolid (3) (Figure S1 and Table S1 in Supplementary Material). These 30 individuals provided us data for 54 posreproductive periods (Table S1 in Supp. Mat.). The individuals were, at least, two years old (del Hoyo et al., 1994) in their first period considered (Euring codes 5 and 6). All individuals reproduced successfully in Spain. The breeding season in the Hen Harrier lasts from March to July (Garc\u0026iacute;a-Mac\u0026iacute;a et al., \u003cem\u003eunpublised report\u003c/em\u003e) so we analysed the movements in the posreproductive periods, which were generally from August to February, although the dates of the individual reproductive periods were accounted for (Table S1 in Supp. Mat.). Where exact dates were not known, the mean breeding dates of this species were considered (from March 21\u003csup\u003est\u003c/sup\u003e to July 4\u003csup\u003eth\u003c/sup\u003e; Garc\u0026iacute;a-Mac\u0026iacute;a et al., \u003cem\u003eunpublised report\u003c/em\u003e).\u003c/p\u003e\n\u003cp\u003eBetween 2019 and 2022, the Hen Harriers were lured with a Eurasian Eagle-Owl and captured with dho-gaza nets. During the marking process, individuals were ringed and morphometric measurements were taken. Sexing was based on the clear sexual colour dimorphism of the species (Ferguson-Lees and Christie, 2001). Individuals were fitted with satellite tracking transmitters (9g Milsar GSMTag-U9 or 10g-12g Ornitela OrniTrack 10/E10 3G GPS/GSM), back-mounted using thoracic Teflon harnesses. Following established guidelines, the weight of the backpacks was less than 3% of the body weight of each individual (Kenward, 2001). The handling time until release of captured individuals was less than one hour to avoid stressing the animals. The devices were set to collect locations every 15\u0026ndash;30 min, depending on battery voltage, season, and time of day. Data packages were transmitted via the digital cellular network. Then the data were filtered with a 30 min period to standardize the sample.\u003c/p\u003e\n\u003cp\u003eCapture of individuals was carried out by expert licenced ringers from the Spanish Ornithological Society (SEO/BirdLife), complying with ethical standards in the capture and tagging of Hen Harriers. The location data of all individuals were provided by SEO/Birdlife and Natural England to the Vertebrate Zoology Research Group of the University of Alicante.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eStrategies classification\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eBefore conducting the analyses, we undertook a qualitative classification of the posreproductive periods of the individuals (Figures 1 and 2). This classification was based on previous studies and reviews (Garc\u0026iacute;a-Mac\u0026iacute;a et al., 2022c; Newton, 2010), and these strategies can be described as:\u0026nbsp;\u003c/p\u003e\n\u003cul\u003e\n \u003cli\u003e\u003cem\u003eMigration\u0026nbsp;\u003c/em\u003e(M): Individuals exhibiting two distinct latitudinal movements (in spring and autumn) during the posreproductive period between breeding and wintering areas. Where the latitudinal distance exceeds 400 km (Kerlinger 1989: Moroll\u0026oacute;n et al., 2024).\u003c/li\u003e\n \u003cli\u003e\u003cem\u003eSedentarism\u003c/em\u003e: Individuals that did not perform two clear latitudinal movements in the posreproductive period and remain in the breeding area.\u003cul style=\"list-style-type: circle;\"\u003e\n \u003cli\u003e\u003cem\u003eSedentarism sensu stricto\u0026nbsp;\u003c/em\u003e(S): Individuals that remained very close to the nest for the entire year, without significant displacements. They only performed routine day-to-day movements throughout the year.\u0026nbsp;\u003c/li\u003e\n \u003cli\u003e\u003cem\u003eSedentarism with posreproductive movements\u0026nbsp;\u003c/em\u003e(SM; or dispersive migration): Individuals which performed all-direction movements far away from their nests after breeding season, and then a return movement toward the starting point in time for the next breeding season. It involved both wandering movements and settling in temporary settlement areas (TSAs). These movements have a component more exploratory and dispersive, as is shown in previous works (Rosier et al., 2006; Moss and Hipkiss, 2014; Wheat et al., 2017; Hegg\u0026oslash;y et al., 2021), similarly to the dispersal movements of immature birds (\u0026lt;2 years old).\u003c/li\u003e\n \u003c/ul\u003e\n \u003c/li\u003e\n\u003c/ul\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eSpatial parameters\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eBased on previous studies on posreproductive movements (McCaslin et al., 2020; Garc\u0026iacute;a-Mac\u0026iacute;a et al., 2022c) two variables were considered to analyse the movements of each period: maximum distance from the reproductive centroid (km; MaxD), that is, the Euclidean distance of the farthest location from the reproductive centroid and the mean Euclidean distance from the reproductive centroid (km; MeanD). The centroid of the breeding area was calculated using the smallest weekly 50% kernel of each individual each year. This smallest kernel indicates the time at which the member of the pair were incubating eggs, as we can see in other species of raptors (L\u0026oacute;pez-L\u0026oacute;pez et al., 2021; Moroll\u0026oacute;n et al., 2022; Garc\u0026iacute;a-Mac\u0026iacute;a et al., 2023) and therefore the locations are focused on the nest of each pair. To calculate these variables, we used the following R packages: \u003cem\u003eadehabitatHR\u003c/em\u003e (Calenge, 2006) to obtain the smallest breeding area, the package \u003cem\u003ergeos\u003c/em\u003e (Bivand and Rundel, 2021) to calculate the breeding centroid and the base R package \u003cem\u003estats\u003c/em\u003e (R Core Team, 2021) to compute the distances during the posreproductive period from the breeding centroid.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eStatistical analysis\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe used a framework of Linear Mixed Models (LMMs) to check the differences between these three strategies (M, S and SM) and sexes. In both models, \u003cem\u003eSex\u003c/em\u003e and \u003cem\u003eStrategy\u003c/em\u003e were fixed effects, while \u003cem\u003eID\u003c/em\u003e and \u003cem\u003ePeriod\u003c/em\u003e were considered as random effects. The response variables were the \u003cem\u003eMaximum distance to the breeding centroid\u003c/em\u003e and the \u003cem\u003eMean distance to the breeding centroid\u003c/em\u003e, with both variables square root transformed to improve model fit. In order to evaluate the significance of mixed effects models, ANOVA tests with Kenward-Roger approximations were performed. Finally, pairwise comparisons using Estimated Marginal Means test (EMM) with Kenward-Roger approximations and with Tukey test method to p-value adjustment were used to find if there are some differences between strategies. Then, we performed a Fisher\u0026rsquo;s exact test (non-parametric test) to analyse the sex distribution among the three different strategies. Finally, we computed a linear regression between the MaxD and the breeding centroid latitude to know if there are any correlation between the position in the Iberian Peninsula of the breeding area and the movements of the Hen Harriers.\u003c/p\u003e\n\u003cp\u003eStatistical analyses were performed with R software version 4.2.1 (R Core Team, 2021), and the level of significance was set at P\u0026lt;0.05. We used the following R packages: LMMs were computed using \u003cem\u003elme4\u0026nbsp;\u003c/em\u003e(Bates et al., 2015), ANOVA was calculated with \u003cem\u003ecar\u0026nbsp;\u003c/em\u003e(Fox and Weisberg, 2019), and the pairwise comparison EMM test with \u003cem\u003eemmeans\u0026nbsp;\u003c/em\u003e(Lenth et al., 2022). Linear regression and Pearson\u0026rsquo;s coefficient were calculated with the package \u003cem\u003eggpubr\u003c/em\u003e (Kassambara, 2020). QGIS software version 3.22.6 (QGIS Development Team, 2022) was used to visualize the breeding areas, centroids and distances of each individual, as well as for the creation of the maps.\u0026nbsp;\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cstrong\u003e\u003cem\u003eStrategies classification\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe 54 posreproductive periods (36 from 18 females and 18 from 12 males) were classified into three strategies: Migration (M), Sedentarism (S) and Sedentarism with posreproductive movements (SM) (Figure 3). The most common strategy was SM, with 37 periods (68.5%), 26 females and 5 males; followed by M with 12 periods (22.2%), 8 females and 4 males; and finally, S with only 5 periods (9.3%), 2 females and 3 males (Table 1).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eSpatial parameters\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe S strategy had the lower values in both variables, MaxD and MeanD. On the contrary, M had the higher values, and in between these two limits we found the SM (Table 1). We found significant differences in ANOVA test in both variables when we compared the different strategies (MeanD: F=12.22, p\u0026lt;0.001; MaxD: F=13.90, p\u0026lt;0.001; Table 2). In addition, the EMM \u003cem\u003epost-hoc\u003c/em\u003e analysis showed these differences in all comparisons except when comparing SDM and SM (Table 3). When we compared both variables between sexes, we observed that there were no differences in MeanD (F=0.498, p=0.488) neither in MaxD (F\u0026lt;0.001, p=0.976) (Table 2). Also were no differences when we compared both Sex and Strategy together MeanD (F=0.15, p=0.870) neither in MaxD (F=0.047, p=0.954)\u003c/p\u003e\n\u003cp\u003eThere were no differences in the distribution between Sexes in the different Strategies in the Fisher\u0026rsquo;s exact test (p\u0026gt;0.05 in all strategies; Figure 4).\u003c/p\u003e\n\u003cp\u003eThe linear regressions in the different strategies between the MaxD and the breeding centroid latitude were not significative (all p\u0026gt;0.05), however there were trends in the strategies; migration strategy (M) showed a positive Pearson\u0026rsquo;s coefficient between both variables (R=0.413), while the sedentary strategies S and SM showed a negative relationship (R=-0.022, R=-0.263, respectively) (Figure 5).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eFor the first time, the posreproductive strategies of the Spanish population of the Hen Harrier have been studied and defined. The analyses found behaviour that exhibits three different posreproductive strategies: migration with a latitudinal movement of more than 400 km; sedentarism \u003cem\u003esensu stricto\u003c/em\u003e; and sedentarism with posreproductive movements or also called dispersive migration (Newton, \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). These patterns are the same to those found in the Red Kite in previous studies (Garc\u0026iacute;a-Mac\u0026iacute;a et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2022c\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe most common strategy in Red Kites was sedentarism with 75% of posreproductive periods while migrants accounted for only 10% (Garc\u0026iacute;a-Mac\u0026iacute;a et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2022c\u003c/span\u003e). In Hen Harriers, the most common strategy was sedentary with posreproductive movements with 68.5% and the least common was sedentary with only 9.3% of posreproductive periods. This result is novel as it was previously believed that the vast majority of the Hen Harrier population in Spain did not perform large movements and was sedentary (del Hoyo et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Garc\u0026iacute;a and Arroyo, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Redpath et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Arroyo, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOne explanation for these patterns could be that in addition to the Common Vole (\u003cem\u003eMicrotus arvalis\u003c/em\u003e), Hen Harriers during the breeding season often feed on chicks and fledglings of other birds before they reach full flight capacity (del Hoyo et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Redalph et al., 2001; Arroyo and Redpath, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Ludwig et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The young of many open-nesting species have a prolonged period of following their parents without reaching full fledging ability, unlike forest species. The lack of this easy resource after the breeding season may lead to a different hunting behaviour, which motivates the search for new food sources, preferably rodents, which is a more dispersed and difficult resource, and consequently a greater dispersal.\u003c/p\u003e \u003cp\u003eThe results show variability in the behaviour of individuals, which may be a logical adaptation to different environmental, trophic and habitat conditions. For example, harriers fly through areas of recently harvested or unharvested cereal or scrubland and this implies different prey that may also vary, such as population explosions of Common Voles or young fledglings. This adaptive plasticity occurs even in the same individuals that change their strategy from one year to the next (As occurs with individual 200452, which remains sedentary during the first two years and in the third year undertakes a migration of almost 950 kilometres to Morocco; see Table S1 in Supp. Mat.), as is also the case of the Red Kite in the Iberian Peninsula (Garc\u0026iacute;a-Mac\u0026iacute;a et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2022c\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOur results also showed that migration strategies have a connection between distance travelled and latitude of the breeding area, as is the case with partial (in lower latitudes) and full migrations (in higher latitudes) throughout Europe. Most of the breeding areas of Hen Harriers that show shorter movements are located further south than those with larger movements. This may be due to the fact that the birds that breed in the Iberian Peninsula do not have as many ranges of movement compared to those that breed in northern Europe or America, which are characterised by longer migrations.\u003c/p\u003e \u003cp\u003eFrom an evolutionary perspective, the behaviour of the Hen Harrier in the Iberian Peninsula can be interpreted as the result of selective pressures specific to the edge of its geographical range. These pressures tend to favour flexibility in movement patterns (Brown, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Brown and Lomolino, \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1998\u003c/span\u003e), enabling individuals to optimise their balance between site fidelity for breeding, resource acquisition during winter, and the reduction of migratory costs. Full migration could be costly in terms of energy and risks associated with movement, whereas complete sedentarism could increase local competition or the risk of food shortages.\u003c/p\u003e \u003cp\u003eThe behavioural flexibility observed in the Hen Harrier in Spain might represent conditions similar to the origins of migratory behaviour (Salewski and Bruderer, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). The increasingly benign climatic conditions of the Iberian Peninsula due to the global warming, its position at the southern limit of the species' range, and other factors, enable the coexistence of sedentary, migratory, and dispersive individuals. This variability provides a 'natural laboratory' where selection can act, favouring behaviours most adaptive to the species as selective pressures shift (Berthold, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Winkler, \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Salewski and Bruderer, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe most widely accepted hypothesis about the origin of migration suggests that this behaviour evolved gradually from local or dispersive movements, which over time increased in distance and complexity (Salewski and Bruderer, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). In this context, the characteristics observed in the Hen Harrier may reflect early stages in the evolution of migration or, alternatively, a partial reversion towards residency in response to favourable environmental conditions. In both cases, natural selection seems to be shaping a diversity of movement strategies, ranging from full migration to more flexible and complex behaviours.\u003c/p\u003e \u003cp\u003eSite fidelity for breeding is a common trait among all individuals in this population, suggesting that this characteristic may have a strong genetic basis or be favoured for its role in reproductive success (Pulido et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Pulido and Berthold, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). However, the variability in migratory behaviour raises questions about the degree of genetic inheritance of these strategies. Although we lack data on kinship relationships to assess whether sedentary individuals originate from sedentary parents or if migrators have migratory progenitors, studies on other species, such as the Red Kite, have shown that these behaviours can change even within the same generation in response to environmental conditions (Garc\u0026iacute;a-Mac\u0026iacute;a et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2022c\u003c/span\u003e), as observed in the Hen Harrier with ID 200452.\u003c/p\u003e \u003cp\u003eThis case highlights how selection can act rapidly to adjust the behaviour of a population to local conditions. If climatic conditions remain favourable, migratory behaviours may gradually be replaced by sedentary strategies, which are considered more energy-efficient. However, intermediate or hybrid behaviours, such as dispersive migration observed in the Hen Harrier, could also be selected between these extremes.\u003c/p\u003e \u003cp\u003eIn conclusion, this example provides insight into the natural selection processes shaping the evolution of migratory behaviour in birds. The coexistence of multiple strategies within a single population underscores the importance of plasticity and variability as key factors in adapting to dynamic environments. As environmental conditions continue to change, these selective processes will persist in defining the optimal behaviour for the population. We propose that such conditions are those required for the emergence or evolutionary selection of migratory or sedentary behaviours within a population, as may have occurred in other species during other periods in range-limit areas.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAnimal Ethics Statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis study did not involve experimental procedures on animals and therefore, according to Spanish legislation, did not require approval from an animal ethics committee. The research involved only the capture and tagging of wild Hen Harriers (Circus cyaneus) with lightweight GPS devices, which is regulated by regional wildlife authorities in Spain. All trapping and marking activities were conducted under permits issued by the competent regional administrations:\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026bull; Departamento de Industria, Transici\u0026oacute;n Energ\u0026eacute;tica y Sostenibilidad, Gobierno Vasco\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026bull; Consejer\u0026iacute;a de Administraci\u0026oacute;n Auton\u0026oacute;mica, Medio Ambiente y Cambio Clim\u0026aacute;tico, Principado de Asturias\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026bull; Consejer\u0026iacute;a de Fomento, Ordenaci\u0026oacute;n del Territorio y Medio Ambiente, Gobierno de Cantabria\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026bull; Consejer\u0026iacute;a de Agricultura, Ganader\u0026iacute;a y Medio Ambiente, Gobierno de La Rioja\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026bull; Departamento de Desarrollo Rural y Medio Ambiente, Gobierno de Navarra (Permit No. 0001-0261-2020-000027)\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026bull; Departamento de Acci\u0026oacute;n Clim\u0026aacute;tica, Alimentaci\u0026oacute;n y Agenda Rural, Generalitat de Catalu\u0026ntilde;a\u003c/p\u003e\n\u003cp\u003e\u0026bull; Consejer\u0026iacute;a de Medio Ambiente, Licencias de Caza y Pesca, Comunidad de Madrid \u0026bull; Consejer\u0026iacute;a de Fomento y Medio Ambiente, Junta de Castilla y Le\u0026oacute;n (Permit No. AUES/CYL/28/2021)\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026bull; Consejer\u0026iacute;a de Agricultura, Ganader\u0026iacute;a y Desarrollo Rural, Gobierno de Castilla-La Mancha Captures were carried out by expert licensed ringers from the Spanish Ornithological Society (SEO/BirdLife), following strict ethical standards for the capture and tagging of wild raptors.\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe results provided in this publication were obtained thanks to the work of SEO/BirdLife in the Migra program and its funding by Fundaci\u0026oacute;n Iberdrola Espa\u0026ntilde;a and Naturand England. \u0026nbsp;Numerous individuals, ornithological groups, ringing groups, research centers, forestry agents and technicians from the autonomous communities were involved. Without their collaboration it would not have been possible to carry out part of the work.\u003c/p\u003e\n\u003cp\u003eMany people from SEO/BirdLife also took part in the work, especially Javier de la Puente, Ana Bermejo and Carmen Fern\u0026aacute;ndez. And of course the staff who helped with other administrative and logistical elements of project management. From Natural England we would especially like to thank for their great collaboration in the capture of individuals to Javier de la Puente, Arturo Rodr\u0026iacute;guez and Juan Mart\u0026iacute;nez.\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was supported by\u0026nbsp;Fundación Iberdrola España (MIGRA program of SEO/BirdLife) and Natural England.\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData Availability Statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll data used in this study are publicly available upon request to data managers in the online data repository Movebank (www.movebank.org). The projects are: \u0026ldquo;Hen Harrier in Spain \u0026ndash; Migra Program in Spain\u0026rdquo; (project ID: 1169461367) and \u0026ldquo;Hen Harriers in Spain\u0026rdquo; (project ID: 1135251191).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors\u0026rsquo; contribution\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eS.M., and V.U. conceived the ideas and designed the methodology, S.L. managed the data. S.M. and J.G.M. analysed the data. S.M. and V.U. wrote the manuscript. S.M., JGM., S.L. and V.U and contributed critically to the drafts. S.L. and V.U. gave final approval for publication. \u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that no conflict of interest exists.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAmar, A. and Redpath, S.M. (2005). 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Fitting Linear Mixed-Effects Models Using lme4. \u003cem\u003eJournal of Statistical Software\u003c/em\u003e. \u003cstrong\u003e67\u003c/strong\u003e(1): 1-48. \u003c/li\u003e\n\u003cli\u003eBerthold P (1999) A comprehensive theory for the evolution, control and adaptability of avian migration. Ostrich 70:1\u0026ndash;11\u003c/li\u003e\n\u003cli\u003eBivand, R. and Rundel, C. (2021). \u003cem\u003ergeos\u003c/em\u003e: Interface to Geometry Engine - Open Source (\u0026apos;GEOS\u0026apos;). R package version 0.5-9. https://CRAN.R-project.org/package=rgeos\u003c/li\u003e\n\u003cli\u003eBrown, J.H. (1984). On the relationship between abundance and distribution of species. \u003cem\u003eThe American Naturalist. \u003c/em\u003e\u003cstrong\u003e124\u003c/strong\u003e(2): 255-279.\u003c/li\u003e\n\u003cli\u003eBrown, J.H. and Lomolino, M.V. (1998). \u003cem\u003eBiogeography.\u003c/em\u003e Sinauer Associates, Inc. Publishers: Sunderland, UK.\u003c/li\u003e\n\u003cli\u003eCalenge, C. 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Thousand Oaks CA: Sage. URL: https://socialsciences.mcmaster.ca/jfox/Books/Companion/\u003c/li\u003e\n\u003cli\u003eGarc\u0026iacute;a, J.T. and Arroyo B.E. (2001). Effect of abiotic factors on reproduction in the centre and periphery of breeding ranges: a comparative analysis in sympatric harriers. \u003cem\u003eEcography\u003c/em\u003e, \u003cem\u003e24\u003c/em\u003e, 393-402.\u003c/li\u003e\n\u003cli\u003eGarc\u0026iacute;a-Mac\u0026iacute;a, J., \u0026Aacute;lvarez, E., Gal\u0026aacute;n, M., Iglesias-Lebrija, J.J., G\u0026aacute;lvez, M., Plana, G., Vallverd\u0026uacute;, N. and Urios, V. (2023). Home range variability and philopatry in Cinereous vultures (\u003cem\u003eAegypius monachus\u003c/em\u003e) breeding in Iberia. \u003cem\u003eAvian Research\u003c/em\u003e. \u003cstrong\u003e14\u003c/strong\u003e: 100134.\u003c/li\u003e\n\u003cli\u003eGarc\u0026iacute;a-Mac\u0026iacute;a, J., De La Puente, J., Bermejo-Bermejo, A., Raab, R., and Urios, V. (2022a). High Variability and Dual Strategy in the Wintering Red Kites (\u003cem\u003eMilvus milvus\u003c/em\u003e). \u003cem\u003eDiversity\u003c/em\u003e. \u003cstrong\u003e14\u003c/strong\u003e(2): 117. \u003c/li\u003e\n\u003cli\u003eGarc\u0026iacute;a-Mac\u0026iacute;a, J., L\u0026oacute;pez-Poveda, G., De La Puente, J., Bermejo-Bermejo, A., Gal\u0026aacute;n, M., \u0026Aacute;lvarez, E., Moroll\u0026oacute;n, S. and Urios, V. (2022b). The variability of juvenile dispersal in an opportunistic raptor. \u003cem\u003eCurrent Zoology\u003c/em\u003e. zoac039, \u003cstrong\u003e69\u003c/strong\u003e(3): 244-254. \u003c/li\u003e\n\u003cli\u003eGarc\u0026iacute;a-Mac\u0026iacute;a, J., Maeso, S., Moroll\u0026oacute;n, S., Simon, L. and Urios, V. Spatial ecology of the Hen Harrier (\u003cem\u003eCircus cyaneus\u003c/em\u003e) during the breeding period in Spain. \u003cem\u003eUnpublised report.\u003c/em\u003e\u003c/li\u003e\n\u003cli\u003eGarc\u0026iacute;a-Mac\u0026iacute;a, J., Pomares, A., De la Puente, J., Bermejo, A., Mart\u0026iacute;nez, J., \u0026Aacute;lvarez, E., Moroll\u0026oacute;n, S. and Urios, V. (2022c). Striking Variability in the Posreproductive Movements of Spanish Red Kites (\u003cem\u003eMilvus milvus\u003c/em\u003e): Three Strategies, Sex Differences, and Changes over Time. \u003cem\u003eAnimals\u003c/em\u003e. \u003cstrong\u003e12\u003c/strong\u003e(21): 2930.\u003c/li\u003e\n\u003cli\u003eGarc\u0026iacute;a-Mac\u0026iacute;a, J., Vidal-Mateo, J., De La Puente, J., Bermejo, A., Raab, R., and Urios, V. (2021). Seasonal differences in migration strategies of Red Kites (\u003cem\u003eMilvus milvus\u003c/em\u003e) wintering in Spain. \u003cem\u003eJournal of Ornithology\u003c/em\u003e. \u003cstrong\u003e163\u003c/strong\u003e(1): 27-36.\u003c/li\u003e\n\u003cli\u003eGarc\u0026iacute;a-Mac\u0026iacute;a, J., Vidal-Mateo, J., de la Puente, J., Bermejo, A., and Urios, V. (2022d). Spatial ecology of the Red Kite (\u003cem\u003eMilvus milvus\u003c/em\u003e) during the breeding period in Spain. \u003cem\u003eOrnis Fennica\u003c/em\u003e. \u003cstrong\u003e99\u003c/strong\u003e: 150-162.\u003c/li\u003e\n\u003cli\u003eHegg\u0026oslash;y, O., Aarvak, T., Ranke, P.S., Solheim, R., \u0026Oslash;ien, I.J. (2021). Home Range and Excursive Post-breeding Movements of Eurasian Eagle-Owls Revealed by GPS Satellite Transmitters. \u003cem\u003eJournal of Raptor Research. \u003c/em\u003e\u003cstrong\u003e55\u003c/strong\u003e: 619-626. \u003c/li\u003e\n\u003cli\u003eKassambara, A. (2020). ggpubr: \u0026apos;ggplot2\u0026apos; Based Publication Ready Plots. R package version 0.4.0. https://CRAN.R-project.org/package=ggpubr\u003c/li\u003e\n\u003cli\u003eKenward, R.E. (2001). \u003cem\u003eA Manual for Wildlife Radio Tagging\u003c/em\u003e. Academic Press: London, UK.\u003c/li\u003e\n\u003cli\u003eKerlinger, P. (1989). \u003cem\u003eFlight strategies of migrating hawks\u003c/em\u003e. Chicago University Press: Chicago, USA. \u003c/li\u003e\n\u003cli\u003eLenth, R. V., Buerkner, P., Herve, M., Love, J., Miguez, F., Riebl, H., and Singmann, H. (2022). Package \u0026ldquo;Emmeans\u0026rdquo;(Version R Package 1.7.2): Estimated Marginal Means, Aka Least-Squares Means. https://CRAN.R-project.org/package=emmeans\u003c/li\u003e\n\u003cli\u003eLimi\u0026ntilde;ana, R., Soutullo, \u0026Aacute;.A, Urios, V. and Reig-Ferrer, A. 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Diversionary feeding and nestling diet of Hen Harriers Circus cyaneus. \u003cem\u003eBird Study\u003c/em\u003e. \u003cstrong\u003e65\u003c/strong\u003e(4): 431-443.\u003c/li\u003e\n\u003cli\u003eMcCaslin, H.M., Caughlin, T.T. and Heath, J.A. (2020). Long-distance natal dispersal is relatively frequent and correlated with environmental factors in a widespread raptor. \u003cem\u003eJournal of Animal Ecology.\u003c/em\u003e \u003cstrong\u003e89\u003c/strong\u003e: 2077-2088. \u003c/li\u003e\n\u003cli\u003eMoroll\u0026oacute;n, S., Garc\u0026iacute;a-Mac\u0026iacute;a, J., Onrubia, A., Lee, S. and Urios, V. (2024) Migration patterns of the breeding Hen Harrier \u003cem\u003eCircus cyaneus\u003c/em\u003e in Spain. \u003cem\u003eBird Study. \u003c/em\u003e\u003cstrong\u003e71\u003c/strong\u003e: 40-47.\u003c/li\u003e\n\u003cli\u003eMoroll\u0026oacute;n, S., Urios, V. and L\u0026oacute;pez-L\u0026oacute;pez, P. (2022). Home-Range Size and Space Use of Territorial Bonelli\u0026rsquo;s Eagles (\u003cem\u003eAquila fasciata\u003c/em\u003e) Tracked by High-Resolution GPS/GSM Telemetry. \u003cem\u003eDiversity\u003c/em\u003e. \u003cstrong\u003e14\u003c/strong\u003e(12): 1082. \u003c/li\u003e\n\u003cli\u003eMoss, E.H.R. and Hipkiss, T. (2014) Home-range size and examples of post-nesting movements for adult golden eagles (Aquila chrysaetos) in boreal Sweden. \u003cem\u003eJournal of Raptor Research\u003c/em\u003e. \u003cstrong\u003e48\u003c/strong\u003e: 93-105.\u003c/li\u003e\n\u003cli\u003eNewton, I. (2010). \u003cem\u003eBird Migration\u003c/em\u003e. William Collins: London, UK. \u003c/li\u003e\n\u003cli\u003ePanuccio, M., Agostini, N., Mellone, U., and Bogliani, G. (2014). Circannual variation in movement patterns of the Black Kite (\u003cem\u003eMilvus migrans migrans\u003c/em\u003e): a review. \u003cem\u003eEthology Ecology and Evolution\u003c/em\u003e. \u003cstrong\u003e26\u003c/strong\u003e(1): 1-18.\u003c/li\u003e\n\u003cli\u003ePulido F, Berthold P (2003) Quantitative genetic analysis of migratory behaviour. In: Berthold P, Gwinner E, Sonnenschein E (eds) Avian migration. Springer, Berlin Heidelberg New York, pp 53\u0026ndash;77\u003c/li\u003e\n\u003cli\u003ePulido F, Berthold P, van Noordwijk AJ (1996) Frequency of migrants and migratory activity are genetically correlated in a bird population: evolutionary implications. Proc Nat Acad Sci USA 93:14642\u0026ndash;14647\u003c/li\u003e\n\u003cli\u003eQGIS Development Team. (2022). QGIS Geographic Information System. Open Source Geospatial Foundation Project. http://www.qgis.org \u003c/li\u003e\n\u003cli\u003eR Core Team (2021). R: A language and environment for statistical computing. RFoundation for Statistical Computing, Vienna, Austria. URL https://www.R-project.org/.\u003c/li\u003e\n\u003cli\u003eRedpath, S.M., Arroyo, B.E., Etheridge, B., Lekie, F.M., Bowman, K. and Thirgood, S. (2002). Temperature and Hen Harrier productivity: from local mechanisms to geographical patterns. \u003cem\u003eEcography.\u003c/em\u003e \u003cstrong\u003e25\u003c/strong\u003e: 533-540. \u003c/li\u003e\n\u003cli\u003eRedpath, S.M., Clarke, R., Madders, M. and Thirgood, S.J. (2001). Assessing raptor diet: comparing pellets, prey remains, and observational data at Hen Harrier nests. \u003cem\u003eCondor.\u003c/em\u003e \u003cstrong\u003e103\u003c/strong\u003e: 184-188.\u003c/li\u003e\n\u003cli\u003eRosier, J.R., Ronan, N.A., Rosenberg, D.K. (2006). Post-breeding Dispersal of Burrowing Owls in an Extensive California Grassland. \u003cem\u003eThe American midland naturalist. \u003c/em\u003e\u003cstrong\u003e155\u003c/strong\u003e(1):\u003cem\u003e \u003c/em\u003e162-167.\u003c/li\u003e\n\u003cli\u003eSalewski, V., \u0026amp; Bruderer, B. (2007). The evolution of bird migration\u0026mdash;a synthesis. \u003cem\u003eNaturwissenschaften\u003c/em\u003e, \u003cem\u003e94\u003c/em\u003e, 268-279.\u003c/li\u003e\n\u003cli\u003eWheat, R.E., Lewis, S.B., Wang, Y., Levi, T. and Wilmers, C.C. (2017). To migrate, stay put, or wander? Varied movement strategies in bald eagles (\u003cem\u003eHaliaeetus leucocephalus\u003c/em\u003e). \u003cem\u003eMovement Ecoogy.\u003c/em\u003e \u003cstrong\u003e5\u003c/strong\u003e: 9. \u003c/li\u003e\n\u003cli\u003eWinkler, D.W. (2005) How do migration and dispersal interact. In: Greenberg R, Marra PP (eds) \u003cem\u003eBirds of two worlds\u003c/em\u003e. Johns Hopkins University Press, Baltimore, 401-413.\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTable 1: Percentages, Mean, Standard Deviation and Range of Maximum and Mean distances to the breeding centroid by strategies.\u003c/p\u003e\n\u003cdiv align=\"\"\u003e\n \u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"706\" style=\"margin-right: calc(12%); width: 88%;\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 23.4407%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eStrategy\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.4345%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSex\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 21.1538%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNo. of posreproductive periods (%)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16.1512%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMean distance to the centroid (km)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 21.6455%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMaximum distance to the centroid (km)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"3\" style=\"width: 23.4407%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMigration\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.4345%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTotal\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 21.1538%;\"\u003e\n \u003cp\u003e\u003cstrong\u003e12 (22.2%)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16.1512%;\"\u003e\n \u003cp\u003e\u003cstrong\u003e318.24 \u0026plusmn; 231.74 (44.1 - 795.52)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 27.7287%;\"\u003e\n \u003cp\u003e\u003cstrong\u003e679.27 \u0026plusmn; 368.7 (120.55 - 1227.91)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 11.4345%;\"\u003e\n \u003cp\u003eFemales\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 21.1538%;\"\u003e\n \u003cp\u003e8 (14.8%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16.1512%;\"\u003e\n \u003cp\u003e292.83 \u0026plusmn; 241.47 (44.1 - 795.52)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 27.7287%;\"\u003e\n \u003cp\u003e673.22 \u0026plusmn; 352.69 (120.55 - 1082.34)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 11.4345%;\"\u003e\n \u003cp\u003eMales\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 21.1538%;\"\u003e\n \u003cp\u003e4 (7.4%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16.1512%;\"\u003e\n \u003cp\u003e369.06 \u0026plusmn; 236 (155.14 - 623.22)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 27.7287%;\"\u003e\n \u003cp\u003e691.38 \u0026plusmn; 455.98 (209.28 - 1227.91)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"3\" style=\"width: 23.4407%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSedentarism \u003cem\u003esensu stricto\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.4345%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTotal\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 21.1538%;\"\u003e\n \u003cp\u003e\u003cstrong\u003e5 (9.3%)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16.1512%;\"\u003e\n \u003cp\u003e\u003cstrong\u003e8.89 \u0026plusmn; 6.02 \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;(3.34 - 19.04)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 27.7287%;\"\u003e\n \u003cp\u003e\u003cstrong\u003e31.94 \u0026plusmn; 24.18 (15.59 - 74.68)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 11.4345%;\"\u003e\n \u003cp\u003eFemales\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 21.1538%;\"\u003e\n \u003cp\u003e2 (3.7%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16.1512%;\"\u003e\n \u003cp\u003e8.2 \u0026plusmn; 0.36 \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;(7.94 - 8.45)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 27.7287%;\"\u003e\n \u003cp\u003e21.89 \u0026plusmn; 1.61 \u0026nbsp; \u0026nbsp;(20.75 - 23.03)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 11.4345%;\"\u003e\n \u003cp\u003eMales\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 21.1538%;\"\u003e\n \u003cp\u003e3 (5.6%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16.1512%;\"\u003e\n \u003cp\u003e9.36 \u0026plusmn; 8.47 \u0026nbsp; \u0026nbsp; \u0026nbsp; (3.34 - 19.04)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 27.7287%;\"\u003e\n \u003cp\u003e38.64 \u0026plusmn; 31.62 (15.59 - 74.68)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"3\" style=\"width: 23.4407%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSedentarism with posreproductive movements or dispersive migration\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.4345%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTotal\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 21.1538%;\"\u003e\n \u003cp\u003e\u003cstrong\u003e37 (68.5%)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16.1512%;\"\u003e\n \u003cp\u003e\u003cstrong\u003e127.54 \u0026plusmn; 78.86 (7.27 - 448.21)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 27.7287%;\"\u003e\n \u003cp\u003e\u003cstrong\u003e254.58 \u0026plusmn; 153.66 (74.95 - 893.34)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 11.4345%;\"\u003e\n \u003cp\u003eFemales\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 21.1538%;\"\u003e\n \u003cp\u003e26 (48.2 %)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16.1512%;\"\u003e\n \u003cp\u003e114.36 \u0026plusmn; 52.15 (20.01 - 235.14)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 27.7287%;\"\u003e\n \u003cp\u003e250.46 \u0026plusmn; 117.5 (101.05 - 560.26)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 11.4345%;\"\u003e\n \u003cp\u003eMales\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 21.1538%;\"\u003e\n \u003cp\u003e11 (20.3%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16.1512%;\"\u003e\n \u003cp\u003e158.71 \u0026plusmn; 118.61 (7.27 - 448.21)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 27.7287%;\"\u003e\n \u003cp\u003e264.33 \u0026plusmn; 224.36 (74.95 - 893.34)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n\u003cp\u003eTable 2: ANOVA test results based on the previous LMMs. df: degrees of freedom, df.res: degrees of freedom of the residuals. Significative differences are shown in bold.\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"537\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 161px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eVariable\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 142px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eFactor\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eF\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e\u003cstrong\u003edf\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53px;\"\u003e\n \u003cp\u003e\u003cstrong\u003egl.res\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 67px;\"\u003e\n \u003cp\u003e\u003cstrong\u003ep-value\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"4\" style=\"width: 161px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMean Distance to Breeding Centroid\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 142px;\"\u003e\n \u003cp\u003e(Intercept)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e85.76\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53px;\"\u003e\n \u003cp\u003e27.09\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 67px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026lt;0.001\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 142px;\"\u003e\n \u003cp\u003eSex\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e0.56\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53px;\"\u003e\n \u003cp\u003e24.41\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 67px;\"\u003e\n \u003cp\u003e0.460\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 142px;\"\u003e\n \u003cp\u003eStrategy\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e12.22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53px;\"\u003e\n \u003cp\u003e34.66\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 67px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026lt;0.001\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 142px;\"\u003e\n \u003cp\u003eSex: Strategy\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e0.15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53px;\"\u003e\n \u003cp\u003e35.64\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 67px;\"\u003e\n \u003cp\u003e0.870\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"4\" style=\"width: 161px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMaximum Distance to Breeding Centroid\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 142px;\"\u003e\n \u003cp\u003e(Intercepto)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e141.60\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53px;\"\u003e\n \u003cp\u003e22.86\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 67px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026lt;0.001\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 142px;\"\u003e\n \u003cp\u003eSex\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53px;\"\u003e\n \u003cp\u003e19.61\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 67px;\"\u003e\n \u003cp\u003e0.974\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 142px;\"\u003e\n \u003cp\u003eStrategy\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e13.90\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53px;\"\u003e\n \u003cp\u003e26.94\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 67px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026lt;0.001\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 142px;\"\u003e\n \u003cp\u003eSex: Strategy\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e0.047\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53px;\"\u003e\n \u003cp\u003e27.79\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 67px;\"\u003e\n \u003cp\u003e0.954\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u0026nbsp;\u003cbr\u003eTable 3: Pairwise comparisons with EMM test of the four strategies. LDM: Large-distance migration, SDM: Short-distance migration, S: Sedentarism, SM: Sedentarism with posreproductive movements. SE: Standard Error, df: degrees of freedom. Significative differences are shown in bold.\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"557\" class=\"fr-table-selection-hover\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 161px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eVariable\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 108px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eContrats\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 81px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eEstimador\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eDE\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e\u003cstrong\u003edf\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 61px;\"\u003e\n \u003cp\u003e\u003cstrong\u003et-ratio\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 63px;\"\u003e\n \u003cp\u003e\u003cstrong\u003ep-value\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"3\" style=\"width: 161px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMean Distance to Breeding Centroid\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 108px;\"\u003e\n \u003cp\u003eM-S\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 81px;\"\u003e\n \u003cp\u003e14.89\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37px;\"\u003e\n \u003cp\u003e2.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e45.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 61px;\"\u003e\n \u003cp\u003e6.06\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 63px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026lt;0.001\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 108px;\"\u003e\n \u003cp\u003eM-SM\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 81px;\"\u003e\n \u003cp\u003e6.96\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e25.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 61px;\"\u003e\n \u003cp\u003e3.56\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 63px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e0.004\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 108px;\"\u003e\n \u003cp\u003eS-SM\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 81px;\"\u003e\n \u003cp\u003e-7.93\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37px;\"\u003e\n \u003cp\u003e2.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e46.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 61px;\"\u003e\n \u003cp\u003e-3.69\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 63px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e0.002\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"3\" style=\"width: 161px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMaximum Distance to Breeding Centroid\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 108px;\"\u003e\n \u003cp\u003eM-S\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 81px;\"\u003e\n \u003cp\u003e19.93\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37px;\"\u003e\n \u003cp\u003e3.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e36.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 61px;\"\u003e\n \u003cp\u003e6.33\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 63px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026lt;0.001\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 108px;\"\u003e\n \u003cp\u003eM-SM\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 81px;\"\u003e\n \u003cp\u003e9.61\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37px;\"\u003e\n \u003cp\u003e2.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e20.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 61px;\"\u003e\n \u003cp\u003e4.63\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 63px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026lt;0.001\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 108px;\"\u003e\n \u003cp\u003eS - SM\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 81px;\"\u003e\n \u003cp\u003e-10.32\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37px;\"\u003e\n \u003cp\u003e2.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 45px;\"\u003e\n \u003cp\u003e36.9 \u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 61px;\"\u003e\n \u003cp\u003e-3.71 \u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 63px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e0.002\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e"}],"fulltextSource":"","fullText":"","funders":[{"identity":"fa1db978-9b7e-4c64-8a76-58461a17767a","identifier":"10.13039/100015942","name":"Fundación Iberdrola España","awardNumber":"0","order_by":0},{"identity":"1a67fcab-044f-4c9f-a876-5b8a9ed85c37","identifier":"10.13039/501100001293","name":"Natural England","awardNumber":"0","order_by":1}],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"University of Alicante","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Animal behaviour, Breeding, Dispersive migration, GPS/GSM, Migration, Raptor, Sedentarism","lastPublishedDoi":"10.21203/rs.3.rs-6445538/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6445538/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eKnowledge about the posreproductive movements of species is important for both conservation and ethology. This study investigates the non-breeding or posreproductive movements of Hen Harriers using GPS/GSM data from 30 individuals over 54 periods from 2019 to 2023. Using maximum and mean distances from the reproductive centroid as behavioural proxy, we classified posreproductive periods into in three strategies: migration (22.2%), sedentarism (9.3%), and sedentarism with posreproductive movements (68.5%). Our results challenge the previous belief that the majority of the Spanish Hen Harrier population was sedentary. The study establishes a relationship between distance travelled and latitude of breeding areas, suggesting environmental adaptation. The Spanish Hen Harrier population, being at the southern limit of its breeding range, exhibits less pronounced differences between sexes or strategies. Raptors with extensive migratory and post-breeding periods are prone to be more vulnerable due to the shortening of the area available for posreproductive movements.\u003c/p\u003e","manuscriptTitle":"Variability in spatial behaviour and migratory strategies selection in a global warming context: the case of the Hen Harrier at the southern limit of its distribution","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-05-05 06:28:28","doi":"10.21203/rs.3.rs-6445538/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"380b8e14-09ca-45cc-a3a3-f16435fe3f1d","owner":[],"postedDate":"May 5th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":47449873,"name":"Animal Science"},{"id":47449874,"name":"Behavioral Ecology"},{"id":47449875,"name":"Animal Behavior"}],"tags":[],"updatedAt":"2025-05-05T06:28:28+00:00","versionOfRecord":[],"versionCreatedAt":"2025-05-05 06:28:28","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6445538","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6445538","identity":"rs-6445538","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}