Variation of migration routes in the Central Asian-breeding Cuculus canorus population influenced by the Qinghai-Tibet Plateau

Variation of migration routes in the Central Asian-breeding Cuculus canorus population influenced by the Qinghai-Tibet Plateau | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Variation of migration routes in the Central Asian-breeding Cuculus canorus population influenced by the Qinghai-Tibet Plateau Zhichang Cheng, Donghui Ma, Lingwang Kong, Mengjie Lu, Xingnan Du, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3967127/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Seasonal climate changes and fluctuations in food resources drive the migratory behavior of birds between their breeding and wintering sites. Migratory birds change their migration routes in response to climate and environmental stresses. Existing studies have indicated that the widely distributed Eurasian Common cuckoo migrates individually during the night, necessitating further research to elucidate its global migration routes. From 2018 to 2021, the migration routes of 13 adult Common cuckoos breeding in the north of the Qinghai-Tibet Plateau were tracked using satellite trackers. We found that (1) the migration routes of this Common cuckoo population mainly separate into three directions: 8 individuals followed the western routes along the western edge of the Qinghai-Tibet Plateau to the south, 3 individuals traversed the eastern routes along the eastern part of the Qinghai-Tibet Plateau and the rest 2 opted for the middle routes across most of the Qinghai-Tibet Plateau to the south. (2) Individual marked with the identifier 201907 exhibited a consistent stopover site selection during autumn migration. However, it changed the breeding area after spring migration. (3) In the context of autumn migration, 4 tracked Common cuckoos (50%) along the western migratory routes exhibited a consistent preference for the Hotan area in Xinjiang as their stopover site. Concurrently, 3 individuals (100%) traversing the eastern migratory route consistently selected stopover locations in the vicinity of Baoshan city, Yunnan. (4) The migration speed of Common cuckoos is significantly higher in spring than in autumn, and the stopover duration of spring migration is significantly lower than that of autumn migration. The daily flight time of spring migration is concentrated between 0–12 clock, while that of autumn migration is between 0–6 clock. (5) Migration routes of the Central Asian Common cuckoo population were limited by the Qinghai-Tibet plateau. The population exhibited high levels of plasticity, with individuals with longer wings tending to select the east and middle routes, while shorter wings were observed more frequently in the west route. Our findings reveal that central Asian common cuckoo populations exhibit a preference for a minimum-time strategy during spring migration, with migration routes selected based on wing length. Cuculus canorus birds migration Qinghai-Tibet Plateau migration routes migrant strategy Figures Figure 1 Figure 2 Figure 3 Figure 4 Background Migration of birds refers to the regular seasonal round-trip movement between breeding and wintering areas. This phenomenon is widespread in nature and widely recognized in the field of taxonomy [1–3]. The primary factors driving avian migration are alterations in climate and shifts in food resources [4–7]. Variations in migrant routes can be attributed to fluctuations in climate as well as the influence of predation pressures [8, 9]. When migratory birds encounter ecological barriers such as mountains, deserts, and oceans, the mortality rate during the migration process significantly increases [10]. To ensure the success of migration, migratory birds opt for favorable meteorological conditions and ensure an ample supply of energy [11, 12]. The growing utilization of satellite tracking for migratory birds has facilitated an ongoing deepening of the study on the selection of migration routes among these avian species. Bird migration is a phenomenon that involves the efficient utilization of temporal and spatial resources. This process serves to alleviate interspecific competition, thereby enhancing breeding success rates and survival. Migration is associated with elevated energy consumption, prompting migratory birds to adopt two primary strategies: time minimum strategy and energy minimum strategy [13–15]. Stopover sites offer migrants locations where they can rest and obtain necessary supplies [16, 17]. Additionally, these sites play a significant role in providing shelter for birds facing adverse weather conditions [18]. The quantity and duration of stopovers directly impact the time it takes for birds to reach their destination, serving as a key determinant in the selection of migratory strategies employed by migrant birds. The flight ability is reflected by wing morphology [19]. Most long-distance migration birds exhibited morphological adaptations such as long wingspan suited for rapid flight [20].Long-distance migrants have a higher wing aspect ratio, longer distal wing and lower wing loading than short-distance migrants or residents [21, 22]. The study will also focus on examining how morphological traits influence the selection of migration routes. The investigation into the migratory behavior of common cuckoos, a paradigmatic interspecific brood parasitic avian species, originated in the 1950s when ornithologists began employing ringing methodologies to gather migratory data for the species [23]. Seel [24] upon consolidating ringing data, identified the common cuckoo as a long-distance migratory bird wintering in the African region. Willemoes et al. [25] utilizing satellite tracking during 2010–2011, followed migratory routes of common cuckoo adults from southern Scandinavia to wintering grounds in Africa. The findings revealed night migration with a narrow-front looped pattern, showing similarities and concentration in migration routes. Juveniles, devoid of direct contact with parents and siblings, migrate independently to their wintering areas guided by their innate migration program [26]. Research on satellite-tracked common cuckoo juveniles during migration indicates that juveniles exhibit faster flight speeds and a later onset of migration in the autumn compared to adults [27]. It is evident that current researches on common cuckoo migration predominantly focuses on the Europe-Africa migratory route, with limited data available on the migratory behavior of the Asian breeding population. Especially concerning the common cuckoo population breeding in Central Asia, whether it still opt for Africa as its wintering range or other areas. In the autumn migration process, encountering significant ecological and geographical barriers may increase migration distance, such as the Qinghai-Tibetan Plateau. How do cuckoos choose their migratory route based on morphological traits? This study focuses on the migratory routes, characteristics, and strategies of the greater common cuckoo's Central Asian breeding population north of the Tibetan Plateau. Method Study area and study population We captured Common cuckoos in the Gansu An’xi Extremely-Arid Desert National Nature Reserve(94° 45' ~ 90° 00' E, 39° 52' ~ 41° 53' N) in the northwest of China which borders the Hami region of Xinjiang to the north and is adjacent to Yumen City and Subei Mongolian Autonomous County to the south [28].The western boundary of the reserve is delineated by the Kumtag Desert, while its southern expanse converges at the junction of the Altun and Qilian Mountain ranges, constituting a paradigmatic region characterized by desert and desertification-prone grasslands. Study area is situated in the Central Asian interior, experiences a continental arid climate characterized by extremely low precipitation and air humidity levels, resulting in high evaporation rates. The annual average precipitation is 50.87 mm, with an annual evaporation of 2381.32 mm. The region is marked by prolonged sunshine duration, significant diurnal temperature variations, and an annual average sunshine duration of 3088 hours. The mean annual temperature is 8.74°C, with extreme highs reaching 42.80°C and extreme lows plummeting to -29.3°C. Strong winds, coupled with frequent sandstorms, prevail, with an average annual wind speed of 2.84 m/s and maximum wind forces reaching 7–9 on the Beaufort scale. The aridity index is 11.7 [29]. Breeding area of this Common cuckoo population occupies the low-lying farmlands and wetland habitats situated on the alluvial fan plain to the south of the Anxi Nature Reserve. The primary host species for its brood parasitism is the Isabelline Shrike ( Lanius isabellinus ), known to nest in the tamarisk ( Tamarix ) groves. Other prevalent species breeding in the same vicinity encompass the Saxaul Sparrow ( Passer ammodendri ), Tree sparrow ( Passer montanus ), Barn Swallow ( Hirundo rustica ), Eurasian Collared Dove ( Streptopelia decaocto ), and Crested Lark ( Galerida cristata ). Filed methods The capture of common cuckoo and the deployment of trackers was conducted annually in June from 2018 to 2021. Within patchy Elaeagnus angustifolia forests, mist nets were strategically set up for capturing other bird species during related research activities. It was during these efforts that Common cuckoos occasionally collided with the mist nets. Prior to releasing the captured Common cuckoos, we measured their physical condition and collected approximately 40 microliters of blood samples. Individuals with a body weight exceeding 100g were equipped with a 4.0g tracking device. In June 2021, mist-netting using Common cuckoo vocalizations as bait was implemented. The tracking devices utilized in this study were HQBG0804 models from Hunan Global Messenger company, with a weight of 4.0g. These trackers employed a combination of BeiDou, GPS, and GLONASS for positioning. The data collected were categorized into four levels of positioning accuracy, namely A, B, C, and D, based on parameters such as Vertical Dilution of Precision (VDOP), Horizontal Dilution of Precision (HDOP), and the number of satellites. For these levels, A accuracy was set at 5 meters, B at 10 meters, C at 20 meters, and D at 100 meters, with a confidence level of 95%. The data acquisition frequency of the satellite trackers was typically set at once per hour, and adjustments could be made in case of low device battery levels. Data were transmitted back through mobile networks, including information on longitude, latitude, instantaneous speed, altitude, temperature, accuracy, heading, and voltage. The gathered blood samples were transported to the laboratory for the individual gender identification of common cuckoos. Gender identification was conducted using primers P2(5′ -TCTGCATCGCTAAATCCTTT- 3′) and P8(5′ -CTCCCAAGGATGAGRAAYTG- 3′) [30]. Data analysis We define the onset of autumn migration for the Common cuckoo as the initiation of their long-distance, uninterrupted flight southward from the breeding sites. The time at which the Common cuckoo reaches the first waypoint before arriving at the wintering sites is designated as the termination of the autumn migration (Wang et al., 2018). The starting of spring migration for the Common cuckoo is defined as the time when the bird departs from the final waypoint before leaving the wintering sites. The termination of spring migration is marked by the time when the Common cuckoo reaches the first waypoint upon arriving at the breeding grounds [31]. During the migration process, if the Common cuckoo stays in a particular area for more than two days, we classify that location as a stopover site [32]. The formula for flight speed is calculated as the distance between consecutive stopover sites divided by the time taken to fly that distance [33]. Due to the data received from stopover sites indicating that the instantaneous speed of the Common cuckoo is below 10 km/h, we determine the duration of stay at each stopover site by calculating the time difference between the first point when significant changes occur in instantaneous speed and location information and the arrival time at that specific stopover site [34]. Migration straightness is calculated as the ratio of Migration Straight-line Distance to the Total Migration Distance [35]. In this study, we employed a linear mixed-effects model to investigate the relationship between the dependent variable ‘migrant route selection’ and several predictor variables. In this process western routes were identified 0, eastern and middle routes were identified 1. The model was fitted using Restricted Maximum Likelihood (REML) and included the fixed effects of ‘tarsus length’, ‘wing length’, and ‘body mass’, as well as a random intercept for the grouping variable ‘sex’. In the Global Messenger software, we exported the satellite-tracked migration route data in .shp format. Subsequently, we added the downloaded imagery layer from Google Earth along with the migration route shp layer to ArcGIS 10.6 to delineate the spring and autumn migration routes of the Common cuckoo. Migration routes of different individuals are depicted in distinct colors, breeding sites are represented by rectangles, and wintering sites are denoted by triangles. Stopover sites are categorized based on the duration of stay, ranging from 2–7 days, 7–14 days, 14–21 days, to more than 21 days, with different-sized circles representing different types of mid-stopover sites. We conducted independent sample t-tests to compare migration parameters among different routes, genders, and seasons. Data for each group are presented as mean ± standard error. The data processing was carried out using R version 4.2.2. Results Migration routes Within the study region, migration routes of 13 Common cuckoos (comprising 7 males and 6 females) were systematically documented from 2018 to 2021 (Table 1 ). Notably, two individuals, identified as 201907 and 202115, achieved successful completion of the autumnal migratory endeavor. Additionally, a singular representative, denoted as 201907, undertook a successful spring migration. Population of common cuckoo breeding in Central Asia manifest three principal autumn migration routes (Fig. 1 ), Specifically, 8 individuals (61.5% of the total) opted for the western route around the Qinghai-Tibet Plateau and the Taklamakan Desert. Notably, 201907 undertakes two consecutive autumn migrations, ultimately wintering in Tanzania. 2 Common cuckoos (15.4% of the total) choosed to traverse the Qilian Mountains from their breeding sites, proceeding southward over the Qinghai-Tibet Plateau via Xining. However, 1 individual lost its signal near Lhasa after reaching that vicinity. 3 common cuckoos (23.1%) selected for the eastern route, circumventing the eastern part of the Qinghai-Tibet Plateau. Notably, Common cuckoo 202115 successfully completed its autumn migration and accomplished wintering in Mozambique (Fig. 1 ). The difference in wintering locations between these two common cuckoos (i.e. identified as 201907, 202115) from the same breeding site implies, to some extent, a relatively weak migratory connectivity within the common cuckoo population. Table 1 Common cuckoo Individuals Tracked from 2018 to 2021 Season Birds ID Tracker ID Gender Departure date Arrival date Migration distance (Km) Tarsus(mm) Wing(mm) Weight(g) Autumn 201801 B008 ♂ 2018.8.3 2268.31 22.25 180 98 201803 B006 ♀ 2018.8.3 2045.23 25.15 215 104.08 201804 B007 ♀ 2018.8.15 3279.48 23.48 200 92.86 201805 B010 ♂ 2018.7.31 1125.68 23.7 219 95.91 201906 B011 ♀ 2019.7.29 6248.16 23.54 205 90.67 201907(1st) B012 ♀ 2019.7.31 2019.11.26 9459.45 24.28 201.2 95.26 201907(2nd) B012 ♀ 2020.7.20 2020.11.19 9840.14 202010 B027 ♀ 2020.7.28 1288.35 21.6 197 92.95 202111 HQP3660 ♂ 2021.7.16 4707.01 23.46 212 110.04 202113 HQP3662 ♂ 2021.7.18 2280.85 22.45 216 101.8 202114 HQP3663 ♀ 2021.7.12 2618.82 25.73 210 126.59 202115 HQP3667 ♂ 2021.8.11 2021.12.7 11594.56 23.61 211 101.7 202116 HQP3669 ♂ 2021.8.16 6748.27 23.31 207 111.78 202117 HQP3671 ♂ 2021.8.9 2139.54 24.24 212 107.16 Spring 201907 B012 ♀ 2020.3.27 2020.5.13 9033.51 202115 HQP3667 ♂ 2022.4.2 9197.32 In the western route of autumn migration, the most utilized stopover site is situated in Hotan, Xinjiang (80.78E, 36.25N), with four Common cuckoos (identified as 201801, 201907, 202116, 202117) choosing this site for resting. On the eastern autumn migration route, the secondary hub for stopovers is Baoding City, Yunnan, China (99.23E, 25.46N), where three Common cuckoos (identified as 202010, 202103, 202115) were observed resting. In the central migration route, both Common cuckoos paused for rest in Delingha City, Qinghai, China (98.26E, 36.62N). Furthermore, Dinder National Park in Sudan (34.91E, 12.23N) represens the stopover site with the longest duration (46.5 days). Following the completion of spring migration, Common cuckoo 201907 did not return to its original breeding site in Guazhou County. Instead, it selected a new breeding site near Barkol Kazakh Autonomous County, Hami City, Xinjiang (91.19–94.48 E, 43.21–45.05N) (Fig. 2 ). During its second autumn migration, this individual chose stopover sites in Pakistan (Mangla, 73.59E, 33.17N), Eritrea (Gash-Barka, 37.56E, 15.27N), and Sudan (Dinder National Park, 34.91E, 12.23N), all of which are characterized by latitudinal proximity (Table 2 ). Table 2 The main stopover sites and durations during the migration process of the Central Asian breeding population of Common cuckoos. Season Stopover site Region Longitude Latitude Birds Arrive time Departure time Duration (day) Autumn Hotan Prefecture China 80.78E 36.25N 201801 2018.8.24 2018.8.29 5.37 201907 2019.8.4 2019.8.10 5.75 202116 2021.8.1 2021.8.13 12.17 202117 2021.8.12 2021.8.20 8.25 Kashi area China 76.49E 38.66N 201804 2018.9.1 2018.9.16 15.12 Delingha China 98.26E 36.62N 201803 2018.8.7 2018.8.18 11.5 201805 2018.8.15 2018.8.25 10.25 Baoshan China 99.23E 25.46N 202010 2020.8.12 20208.31 18.25 202113 2021.8.7 2021.8.27 20.75 202115 2021.8.18 2021.9.9 21.75 Mangla Pakistan 73.59E 33.17N 201907(1st) 2019.8.13 2019.8.27 14.5 201907(2nd) 2020.7.27 2020.8.20 24.5 202116 2021.8.21 2021.9.3 13.42 Pathankot India 75.81E 35.54N 201906 2019.8.14 2019.9.5 22 Gash-Barka Eritrea 37.56E 15.27 201907(1st) 2019.9.14 2019.9.29 14.5 201907(2nd) 2020.9.14 2019.9.26 12.5 Dinder National Park Sudan 34.91E 12.23N 201907(1st) 2019.9.29 2019.11.15 46.5 201907(2nd) 2020.10.11 2020.11.6 26.5 Spring Mambasa Congo 29.70 2.08 201907 2020.4.4 2020.4.18 13.25 Agew Awi Zone Ethiopia 36.29E 10.56N 201907 2020.4.20 2020.4.28 7.5 Sarayan Iran 58.19E 33.18N 201907 2020.5.1 2020.5.7 6.25 Luuq Somalia 42.77E 3,34N 202115 2022.4.21 2021.4.25 4.25 Migrations traits The results indicated that the Autumn migration distance of 201907 for the first and second subjects were 9459.45km and 9840.14km, respectively. However, the Autumn migration distance of 202115 was 11594.56km. We used independent sample t-test to compare Common cuckoo's autumn and spring migration (Fig. 3), a significant difference emerged in the migratory speed. Specifically, the spring migration speed was notably higher than that observed in autumn migration (spring speed: 789.80 ± 314.68 km/day, autumn speed: 548.61 ± 340.37 km/day, P = 0.002). Furthermore, the duration of stopover at intermediate sites during spring migration was significantly shorter compared to autumn migration (spring stopover duration: 9.97 ± 7.16 days, autumn stopover duration 13.28 ± 8.11 days, P = 0.01). However, there was no statistically significant difference in migratory straightness between spring and autumn migrations (spring straightness: 0.88 ± 0.13, autumn straightness: 0.88 ± 0.14, P = 0.21). Statistical analysis of instantaneous speed data from 13 common cuckoos during migration in the period of 2018–2021, excluding data points with instantaneous speeds below 10 km/h, revealed distinct patterns. During autumn migration (Fig. 4 ), Common cuckoo migration predominantly occurred during the night, concentrated from 0:00 to 6:00 (84.77% of the total migration time). In contrast, during spring migration, the daily migration time span was concentrated from 0:00 to 12:00 (89.08%). There was no significant variation in the selection of migration routes based on the tarsus length and body mass of common cuckoos. However, wing length have a significant difference between various migration routes. Wing length of Common cuckoos that select middle and east migration routes is longer than individuals of west migration routes (Table 3 ). Table 3 The impact of migration route selection on the response of morphological traits. linear mixed model parameters Fixed effects β ± SE df t P Intercept -0.893 ± 3.115 9 -0.287 0.781 Tarsus length -0.181 ± 0.149 9 -1.216 0.255 Wing length 0.033 ± 0.014 9 2.311 0.046 Body mass -0.012 ± 0.016 9 -0.765 0.464 Random effects β ± SD n Results of VCA (%) sex 0.001 ± 0.001 13 0.5 Residual 0.199 ± 0.447 13 99.5 SE of fixed effects is the standard error of the mean; SD of random effects is the square root of the variance. Discussion Our findings reveal three primary autumn migration routes in the Central Asian common cuckoo population: the western route initiates along the southern edge of the Taklamakan Desert and the western edge of the Himalayas, the central route traverses the Tibetan Plateau, and the eastern route follows the edge of the Tibetan Plateau. The variation of migration routes suggests a significant impact of the Qinghai-Tibet Plateau, a large ecological barrier, on the common cuckoo, leading to a high variation in their migratory routes. Different species populations may adopt similar strategies when facing same large ecological barriers like the Tibetan Plateau. Juhant and Bildstein (2017) [36] identified four main migration corridors: the western Himalayan corridor, the eastern Himalayan corridor, the north-south corridor, and the corridor traversing the Himalayas in their study on raptor migration through the Tibetan Plateau. Correspondingly, Migration routes of black kites are similar with the west route [37]. These routes encompass traversing the Taklamakan Desert and the Tianshan Mountains before arriving at their breeding sites. Regarding the middle routes of autumn migration is not fully realized, there is a discernible inclination toward crossing, and the potential for common cuckoo to detour to the southern part of the Tibetan Plateau later in the season is not excluded. Lee et al. [38] discovered that the cuckoo population breeding in Korea migrates to Southeast Africa during the winter, exhibiting migration routes and wintering areas like those of the eastern route of the Central Asian population. Conversely, common cuckoos breeding in Beijing follow a route around the southern edge of the Qinghai-Tibetan Plateau, resembling the western route in autumn migration. Consequently, we posited various species or populations, when confronted with the same ecological barrier, exhibit a remarkable convergence in their selection of migratory routes. La Sorte and Fink [39] based on information from the eBird platform, analyzed seven years of migration routes for 55 terrestrial bird species. They found that ecological barriers increase the flexibility of migratory routes for birds, as exemplified by the diverse choices of migration routes among these 55 species across the American continent. Individuals of Common cuckoo on the same migration route choose identical stopover sites which may be associated with environmental factors during the migration process. Thorup et al. [40] observed a propensity for Common cuckoos to select regions characterized by elevated vegetation cover during migration in their investigation of migratory bird dynamics and resource tracking. Another pivotal determinant could be the shared encounter of individuals on the same migratory route with homogeneous ecological impediments. The presence of substantial ecological barriers has the potential to protract the migratory duration for migrant birds engaged in long-distance flights with migration timings and routes demonstrating seasonal differentiations. The existence of such ecological barriers serves to prompt migratory birds to converge upon designated intermediate rest sites for strategic energy storage and replenishment [39]. During its second autumn migration, the selection of the same stopover sites by 201907 was postulated to be linked to its protracted memory retention. Migrant birds engaged in extensive long-distance flights often exhibit an extended duration of memory retention for stopover sites [41–43]. Research in the peregrine populations of the Eurasian Arctic revealed that long-term memory emerged as the most probable selective agent for the divergence observed in ADCY8 [44]. Between different seasons, the speed of spring migration is significantly higher than that of autumn migration, and the duration of stopovers is notably shorter in spring than in autumn. Analyzing data with instantaneous speeds exceeding 10 km/h, we also observed that the common cuckoo exhibits typical nocturnal migratory behavior, with longer daily flight times during spring migration compared to autumn migration. Birds generally prefer migrating with tailwinds [45], as seen in species such as the European Honey Buzzard ( Pernis apivorus ), White Stork ( Ciconia ciconia ), and Western Osprey ( Pandion haliaetus ). The strength of tailwinds is higher in spring than in autumn, providing a plausible explanation for the observed phenomenon of migratory birds exhibiting faster migration speeds during the spring compared to the autumn season [46]. The faster migratory speed and reduced stopover duration observed in the common cuckoo during spring migration suggested the adoption of a time-minimization strategy. This spring migration strategy aligns with that commonly observed in many avian species, wherein the pursuit of a time-minimization approach allows birds to secure high-quality breeding habitats earlier, thus enhancing reproductive success [47, 48]. Additionally, this strategy diminishes the frequency and duration of stopovers during migration, thereby mitigating the risk of predation by avian predators [49]. Given the common cuckoo's status as a brood parasite without nest-building behaviors, the time minimization strategy employed during spring migration may be linked to the migratory tactics of its hosts. Birds of the Shrike family ( Laniidae ) exhibited faster spring migration speeds compared to autumn migration, and their selection of stopover sites tends to favor areas with abundant vegetation. During spring migration, these birds adopt a strategy that minimizes time spent in transit, aiming to reach their breeding grounds quickly and aligning with the peak of vegetation coverage upon arrival [50–52]. Research has indicated common cuckoos tend to initiate its migration approximately a month later than its parasitic host [53]. Therefore, the expedited arrival at breeding grounds during spring migration in common cuckoos may be construed as an adaptive response aligning with the reproductive timing of their hosts. Our results indicate that individuals opting for eastward and middle migration routes exhibit longer wing length compared to those selecting westward routes. Since the eastern and middle routes are longer than the western routes, Individuals choosing the eastern and central routes may possess longer wings to improve flight efficiency, thus facilitating their ability to cover longer migration distances. This difference has been observed in other migrant birds, Eurasian Blackcap ( Sylvia atricapilla ) has been reported to exhibit various migration routes among individuals, a phenomenon attributed to differences in wing length [54]. Hahn et al. [55] found that longer wing provides benefits in adjusting the speed of migrants to their flight and arrival time. Additionally, the habitats along western migration routes mainly consist of deserts and high mountains, while eastern migration routes offer relatively abundant habitats. Therefore, this result suggests a potential trade-off between morphological traits and the availability of better stopover sites. The reason why the eastern and middle migration routes, which involve longer migration distances, are chosen by cuckoos with relatively longer wing length may be that longer wings provide them with higher flight efficiency and better selection of stopover sites along the way. Conclusion Our study revealed that the migration routes of the common cuckoos breeding in Central Asia are influenced by the Qinghai-Tibet Plateau. It demonstrated the migration strategy of the common cuckoo and suggested that their selection of migration routes may be influenced by wing length. Constrained by data limitations, the investigation into the migratory behavior of the common cuckoo population breeding in Central Asia necessitates ongoing efforts. Further substantiation is required for tracking the middle routes during autumn migration. Declarations Availability of data and materials The datasets analysed during the current study are available from the corresponding author on reasonable request. Acknowledgements We thank the management staff of Gansu An’xi Extremely arid Desert National Nature Reserve for allowing and helping us to conduct this study in the area. We also appreciate the support from Hunan Global Messenger company. Funding Financial support of this study was provided by the National Natural Science Foundation of China (Grants 31672296). Ethics approval and consent to participate The study was approved by the Ethics Committee of School of Life Sciences of Lanzhou University (File no 20180330), and strictly complied with the requirements of the Chinese Wild Animal Protection Law. Consent for publication Not applicable. Competing interests The authors declare no conflicts of interest to any other affiliations or persons. Contributions: ZC wrote this manuscript, developed most of the data summary, and conducted the data analysis. DM and LK participated in data analysis and field activities. ML and XD collected the filed data. XB designed the experiments and refined this manuscript. References Alerstam T, Hedenström A, Åkesson S: Long-distance migration: evolution and determinants . Oikos 2003, 103 (2):247–260. Lack D: Bird migration and natural selection . Oikos 1968:1–9. 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Oikos 2016, 125 (10):1496–1507. Buechley ER, Oppel S, Beatty WS, Nikolov SC, Dobrev V, Arkumarev V, Saravia V, Bougain C, Bounas A, Kret E: Identifying critical migratory bottlenecks and high-use areas for an endangered migratory soaring bird across three continents . Journal of Avian Biology 2018, 49 (7):e01629. Mi C, Møller AP, Guo Y: Annual spatio-temporal migration patterns of Hooded Cranes wintering in Izumi based on satellite tracking and their implications for conservation . Avian Research 2018, 9 :1–9. Abrahms B, Seidel DP, Dougherty E, Hazen EL, Bograd SJ, Wilson AM, Weldon McNutt J, Costa DP, Blake S, Brashares JS: Suite of simple metrics reveals common movement syndromes across vertebrate taxa . Movement ecology 2017, 5 (1):1–11. Juhant MA, Bildstein KL: Raptor migration across and around the Himalayas . Bird migration across the Himalayas: wetland functioning amidst mountains and glaciers 2017:98–116. Kumar N, Gupta U, Jhala YV, Qureshi Q, Gosler AG, Sergio F: GPS-telemetry unveils the regular high-elevation crossing of the Himalayas by a migratory raptor: implications for definition of a “Central Asian Flyway” . Scientific Reports 2020, 10 (1):15988. Lee J-W, Kang S-G, Lee J-Y, Kim H-N, Jin S-J, Bae G-W, Hur W-H, Park J-Y: Long-distance migration of Korean common cuckoos with different host specificities . Global Ecology and Conservation 2023, 43 :e02426. La Sorte FA, Fink D: Migration distance, ecological barriers and en-route variation in the migratory behaviour of terrestrial bird populations . Global Ecology and Biogeography 2017, 26 (2):216–227. Thorup K, Tøttrup AP, Willemoes M, Klaassen RH, Strandberg R, Vega ML, Dasari HP, Araújo MB, Wikelski M, Rahbek C: Resource tracking within and across continents in long-distance bird migrants . Science Advances 2017, 3 (1):e1601360. Mettke-Hofmann C, Gwinner E: Long-term memory for a life on the move . Proceedings of the National Academy of Sciences 2003, 100 (10):5863–5866. Paruk JD, Chickering MD, Long IV D, Uher-Koch H, East A, Poleschook D, Gumm V, Hanson W, Adams EM, Kovach KA: Winter site fidelity and winter movements in Common Loons (Gavia immer) across North America . The Condor: Ornithological Applications 2015, 117 (4):485–493. Rappole JH, Jones P: Evolution of old and new world migration systems . Ardea 2003. Gu Z, Pan S, Lin Z, Hu L, Dai X, Chang J, Xue Y, Su H, Long J, Sun M: Climate-driven flyway changes and memory-based long-distance migration . Nature 2021, 591 (7849):259–264. Rus AI, Duerr AE, Miller TA, Belthoff JR, Katzner TE: Counterintuitive roles of experience and weather on migratory performance . The Auk: Ornithological Advances 2017, 134 (3):485–497. Ferretti A, Rattenborg NC, Ruf T, McWilliams SR, Cardinale M, Fusani L: Sleeping unsafely tucked in to conserve energy in a nocturnal migratory songbird . Current Biology 2019, 29 (16):2766–2772. e2764. Bêty J, Giroux J-F, Gauthier G: Individual variation in timing of migration: causes and reproductive consequences in greater snow geese (Anser caerulescens atlanticus) . Behavioral Ecology and Sociobiology 2004, 57 :1–8. Low M, Arlt D, Pärt T, Öberg M: Delayed timing of breeding as a cost of reproduction . Journal of Avian Biology 2015, 46 (4):325–331. Zhijun M, Bo L, Jiakuan C: Study on the utilitizaion of stopover sites and migration strategies of migratory birds . ACTA ECOLOGICA SINICA 2005, 25 (6):9. Macías-Torres P, Alerstam T, Andersson A, Bäckman J, Thorup K, Tøttrup AP, Sjöberg S: Activity patterns throughout the annual cycle in a long-distance migratory songbird, the red-backed shrike Lanius collurio . Movement Ecology 2022, 10 (1):1–13. Papageorgiou D, Barboutis C, Kassara C, Giokas S: Habitat selection of woodchat shrikes Lanius senator during spring stopover is related to foraging strategy . Current Zoology 2017, 63 (2):139–149. Pedersen L, Onrubia A, Vardanis Y, Barboutis C, Waasdorp S, van Helvert M, Geertsma M, Ekberg P, Willemoes M, Strandberg R: Remarkably similar migration patterns between different red-backed shrike populations suggest that migration rather than breeding area phenology determines the annual cycle . Journal of Avian Biology 2020, 51 (10). Zhou B, Liang W: Seasonal increase in nest defense, but not egg rejection, in a cuckoo host . Avian Research 2024, 15 :100154. Ożarowska A, Zaniewicz G, Meissner WJCZ: Sex and age-specific differences in wing pointedness and wing length in blackcaps Sylvia atricapilla migrating through the southern Baltic coast . 2021, 67 (3):271–277. Hahn S, Korner-Nievergelt F, Emmenegger T, Amrhein V, Csörgő T, Gursoy A, Ilieva M, Kverek P, Pérez‐Tris J, Pirrello SJE et al : Longer wings for faster springs–wing length relates to spring phenology in a long‐distance migrant across its range . 2016, 6 (1):68–77. Additional Declarations No competing interests reported. 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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-3967127","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":274477994,"identity":"70adad3f-22d1-42ad-b612-722def45b0f4","order_by":0,"name":"Zhichang Cheng","email":"","orcid":"","institution":"Lanzhou University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Zhichang","middleName":"","lastName":"Cheng","suffix":""},{"id":274477995,"identity":"aa174d58-8510-4007-a860-3d81c6e4f5db","order_by":1,"name":"Donghui Ma","email":"","orcid":"","institution":"Lanzhou University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Donghui","middleName":"","lastName":"Ma","suffix":""},{"id":274477996,"identity":"19d4c078-216f-423b-b4a4-4a8235fcfbf2","order_by":2,"name":"Lingwang Kong","email":"","orcid":"","institution":"Lanzhou University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Lingwang","middleName":"","lastName":"Kong","suffix":""},{"id":274477997,"identity":"167d971a-44c5-4541-bf3d-e2008d7396f5","order_by":3,"name":"Mengjie Lu","email":"","orcid":"","institution":"Lanzhou University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Mengjie","middleName":"","lastName":"Lu","suffix":""},{"id":274477998,"identity":"3950c284-67ef-4760-b95a-b27575521d22","order_by":4,"name":"Xingnan Du","email":"","orcid":"","institution":"Lanzhou University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Xingnan","middleName":"","lastName":"Du","suffix":""},{"id":274477999,"identity":"a0ffb6dc-918e-4c4c-ae18-133b6711ab48","order_by":5,"name":"Xinkang Bao","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAuElEQVRIiWNgGAWjYBACAyBmbKiwAHMkSNByRoJULY1tpGgxZz98THLmPAl7gwPMB2/zMNjlEdRi2ZOWbLhxmwSzwQG2ZGsehuRiwg47kGP48OE2CTaDAzxm0jwMBxIbCGo5/8bg4MM5EjwGB/i/EanlBtCWjQ0SEkBb2IjTYjnjWbLhjGMSBpKH2Ywt5xgkE9Zizp98TLKnxsae73jzwxtvKuwIa0EAZrA7iVc/CkbBKBgFowAPAAAigDckOPC7sAAAAABJRU5ErkJggg==","orcid":"","institution":"Lanzhou University","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Xinkang","middleName":"","lastName":"Bao","suffix":""}],"badges":[],"createdAt":"2024-02-18 14:04:37","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3967127/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3967127/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":51663583,"identity":"59f87b6c-1b9b-485d-974d-ff7e4ea1a6c4","added_by":"auto","created_at":"2024-02-26 20:32:14","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":2388478,"visible":true,"origin":"","legend":"\u003cp\u003eMap Legend for the Autumn Migration Routes of 13 Common cuckoos. Triangles represent wintering areas, squares represent breeding areas, circles of different sizes represent the duration of stopovers at resting sites, and lines of different colors represent the autumn migration routes of different individuals.\u003c/p\u003e","description":"","filename":"Figure1.png","url":"https://assets-eu.researchsquare.com/files/rs-3967127/v1/07fc35184f60ee8de5d0f61a.png"},{"id":51663318,"identity":"717eaf32-927d-4a32-b588-5541209c0f92","added_by":"auto","created_at":"2024-02-26 20:24:14","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":2453539,"visible":true,"origin":"","legend":"\u003cp\u003eMap Legend Refinement for Summary of Spring and Autumn Migration Routes for Common cuckoos 201907 and 202115. Triangles represent wintering areas, squares represent breeding areas, circles of different sizes represent the duration of stopovers at resting sites. Purple and light purple respectively represent the autumn migration route and spring migration route of the cuckoo 202115, while black and gray respectively represent the autumn migration route and spring migration route of the cuckoo 201907.\u003c/p\u003e","description":"","filename":"Figure2.png","url":"https://assets-eu.researchsquare.com/files/rs-3967127/v1/f0d3c539ecd296a8d71af700.png"},{"id":51663315,"identity":"83a1d568-71a7-4ae4-90da-07e982a7dba4","added_by":"auto","created_at":"2024-02-26 20:24:14","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":92450,"visible":true,"origin":"","legend":"\u003cp\u003eStatistical Analysis of Migration Parameters among Different Migration Seasons.\u003c/p\u003e","description":"","filename":"Figure3.png","url":"https://assets-eu.researchsquare.com/files/rs-3967127/v1/5a14a31fb9ec70bda6f54747.png"},{"id":51663317,"identity":"e8613918-fcab-4c7e-9fe8-2742982c7e42","added_by":"auto","created_at":"2024-02-26 20:24:14","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":45548,"visible":true,"origin":"","legend":"\u003cp\u003eStatistical Analysis of Daily Flight Durations during Spring and Autumn Migrations of Common cuckoos\u003c/p\u003e","description":"","filename":"Figure4.png","url":"https://assets-eu.researchsquare.com/files/rs-3967127/v1/ef23d111b46e73b8a2ec1b40.png"},{"id":51874024,"identity":"ee9847f2-9a90-4ea7-b408-f23f42aba434","added_by":"auto","created_at":"2024-03-01 17:45:33","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":5734340,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3967127/v1/b75567ef-82a2-4d98-b5ad-d6529fdf991f.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Variation of migration routes in the Central Asian-breeding Cuculus canorus population influenced by the Qinghai-Tibet Plateau","fulltext":[{"header":"Background","content":"\u003cp\u003eMigration of birds refers to the regular seasonal round-trip movement between breeding and wintering areas. This phenomenon is widespread in nature and widely recognized in the field of taxonomy [1\u0026ndash;3]. The primary factors driving avian migration are alterations in climate and shifts in food resources [4\u0026ndash;7]. Variations in migrant routes can be attributed to fluctuations in climate as well as the influence of predation pressures [8, 9]. When migratory birds encounter ecological barriers such as mountains, deserts, and oceans, the mortality rate during the migration process significantly increases [10]. To ensure the success of migration, migratory birds opt for favorable meteorological conditions and ensure an ample supply of energy [11, 12]. The growing utilization of satellite tracking for migratory birds has facilitated an ongoing deepening of the study on the selection of migration routes among these avian species.\u003c/p\u003e \u003cp\u003eBird migration is a phenomenon that involves the efficient utilization of temporal and spatial resources. This process serves to alleviate interspecific competition, thereby enhancing breeding success rates and survival. Migration is associated with elevated energy consumption, prompting migratory birds to adopt two primary strategies: time minimum strategy and energy minimum strategy [13\u0026ndash;15]. Stopover sites offer migrants locations where they can rest and obtain necessary supplies [16, 17]. Additionally, these sites play a significant role in providing shelter for birds facing adverse weather conditions [18]. The quantity and duration of stopovers directly impact the time it takes for birds to reach their destination, serving as a key determinant in the selection of migratory strategies employed by migrant birds. The flight ability is reflected by wing morphology [19]. Most long-distance migration birds exhibited morphological adaptations such as long wingspan suited for rapid flight [20].Long-distance migrants have a higher wing aspect ratio, longer distal wing and lower wing loading than short-distance migrants or residents [21, 22]. The study will also focus on examining how morphological traits influence the selection of migration routes.\u003c/p\u003e \u003cp\u003eThe investigation into the migratory behavior of common cuckoos, a paradigmatic interspecific brood parasitic avian species, originated in the 1950s when ornithologists began employing ringing methodologies to gather migratory data for the species [23]. Seel [24] upon consolidating ringing data, identified the common cuckoo as a long-distance migratory bird wintering in the African region. Willemoes et al. [25] utilizing satellite tracking during 2010\u0026ndash;2011, followed migratory routes of common cuckoo adults from southern Scandinavia to wintering grounds in Africa. The findings revealed night migration with a narrow-front looped pattern, showing similarities and concentration in migration routes. Juveniles, devoid of direct contact with parents and siblings, migrate independently to their wintering areas guided by their innate migration program [26]. Research on satellite-tracked common cuckoo juveniles during migration indicates that juveniles exhibit faster flight speeds and a later onset of migration in the autumn compared to adults [27]. It is evident that current researches on common cuckoo migration predominantly focuses on the Europe-Africa migratory route, with limited data available on the migratory behavior of the Asian breeding population. Especially concerning the common cuckoo population breeding in Central Asia, whether it still opt for Africa as its wintering range or other areas. In the autumn migration process, encountering significant ecological and geographical barriers may increase migration distance, such as the Qinghai-Tibetan Plateau. How do cuckoos choose their migratory route based on morphological traits? This study focuses on the migratory routes, characteristics, and strategies of the greater common cuckoo's Central Asian breeding population north of the Tibetan Plateau.\u003c/p\u003e"},{"header":"Method","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy area and study population\u003c/h2\u003e \u003cp\u003eWe captured Common cuckoos in the Gansu An\u0026rsquo;xi Extremely-Arid Desert National Nature Reserve(94\u0026deg; 45' ~ 90\u0026deg; 00' E, 39\u0026deg; 52' ~ 41\u0026deg; 53' N) in the northwest of China which borders the Hami region of Xinjiang to the north and is adjacent to Yumen City and Subei Mongolian Autonomous County to the south [28].The western boundary of the reserve is delineated by the Kumtag Desert, while its southern expanse converges at the junction of the Altun and Qilian Mountain ranges, constituting a paradigmatic region characterized by desert and desertification-prone grasslands.\u003c/p\u003e \u003cp\u003eStudy area is situated in the Central Asian interior, experiences a continental arid climate characterized by extremely low precipitation and air humidity levels, resulting in high evaporation rates. The annual average precipitation is 50.87 mm, with an annual evaporation of 2381.32 mm. The region is marked by prolonged sunshine duration, significant diurnal temperature variations, and an annual average sunshine duration of 3088 hours. The mean annual temperature is 8.74\u0026deg;C, with extreme highs reaching 42.80\u0026deg;C and extreme lows plummeting to -29.3\u0026deg;C. Strong winds, coupled with frequent sandstorms, prevail, with an average annual wind speed of 2.84 m/s and maximum wind forces reaching 7\u0026ndash;9 on the Beaufort scale. The aridity index is 11.7 [29].\u003c/p\u003e \u003cp\u003eBreeding area of this Common cuckoo population occupies the low-lying farmlands and wetland habitats situated on the alluvial fan plain to the south of the Anxi Nature Reserve. The primary host species for its brood parasitism is the Isabelline Shrike (\u003cem\u003eLanius isabellinus\u003c/em\u003e), known to nest in the tamarisk (\u003cem\u003eTamarix\u003c/em\u003e) groves. Other prevalent species breeding in the same vicinity encompass the Saxaul Sparrow (\u003cem\u003ePasser ammodendri\u003c/em\u003e), Tree sparrow (\u003cem\u003ePasser montanus\u003c/em\u003e), Barn Swallow (\u003cem\u003eHirundo rustica\u003c/em\u003e), Eurasian Collared Dove (\u003cem\u003eStreptopelia decaocto\u003c/em\u003e), and Crested Lark (\u003cem\u003eGalerida cristata\u003c/em\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eFiled methods\u003c/h2\u003e \u003cp\u003eThe capture of common cuckoo and the deployment of trackers was conducted annually in June from 2018 to 2021. Within patchy Elaeagnus angustifolia forests, mist nets were strategically set up for capturing other bird species during related research activities. It was during these efforts that Common cuckoos occasionally collided with the mist nets. Prior to releasing the captured Common cuckoos, we measured their physical condition and collected approximately 40 microliters of blood samples. Individuals with a body weight exceeding 100g were equipped with a 4.0g tracking device. In June 2021, mist-netting using Common cuckoo vocalizations as bait was implemented.\u003c/p\u003e \u003cp\u003eThe tracking devices utilized in this study were HQBG0804 models from Hunan Global Messenger company, with a weight of 4.0g. These trackers employed a combination of BeiDou, GPS, and GLONASS for positioning. The data collected were categorized into four levels of positioning accuracy, namely A, B, C, and D, based on parameters such as Vertical Dilution of Precision (VDOP), Horizontal Dilution of Precision (HDOP), and the number of satellites. For these levels, A accuracy was set at 5 meters, B at 10 meters, C at 20 meters, and D at 100 meters, with a confidence level of 95%. The data acquisition frequency of the satellite trackers was typically set at once per hour, and adjustments could be made in case of low device battery levels. Data were transmitted back through mobile networks, including information on longitude, latitude, instantaneous speed, altitude, temperature, accuracy, heading, and voltage. The gathered blood samples were transported to the laboratory for the individual gender identification of common cuckoos. Gender identification was conducted using primers P2(5\u0026prime; -TCTGCATCGCTAAATCCTTT- 3\u0026prime;) and P8(5\u0026prime; -CTCCCAAGGATGAGRAAYTG- 3\u0026prime;) [30].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eData analysis\u003c/h2\u003e \u003cp\u003eWe define the onset of autumn migration for the Common cuckoo as the initiation of their long-distance, uninterrupted flight southward from the breeding sites. The time at which the Common cuckoo reaches the first waypoint before arriving at the wintering sites is designated as the termination of the autumn migration (Wang et al., 2018). The starting of spring migration for the Common cuckoo is defined as the time when the bird departs from the final waypoint before leaving the wintering sites. The termination of spring migration is marked by the time when the Common cuckoo reaches the first waypoint upon arriving at the breeding grounds [31]. During the migration process, if the Common cuckoo stays in a particular area for more than two days, we classify that location as a stopover site [32]. The formula for flight speed is calculated as the distance between consecutive stopover sites divided by the time taken to fly that distance [33]. Due to the data received from stopover sites indicating that the instantaneous speed of the Common cuckoo is below 10 km/h, we determine the duration of stay at each stopover site by calculating the time difference between the first point when significant changes occur in instantaneous speed and location information and the arrival time at that specific stopover site [34]. Migration straightness is calculated as the ratio of Migration Straight-line Distance to the Total Migration Distance [35].\u003c/p\u003e \u003cp\u003eIn this study, we employed a linear mixed-effects model to investigate the relationship between the dependent variable \u0026lsquo;migrant route selection\u0026rsquo; and several predictor variables. In this process western routes were identified 0, eastern and middle routes were identified 1. The model was fitted using Restricted Maximum Likelihood (REML) and included the fixed effects of \u0026lsquo;tarsus length\u0026rsquo;, \u0026lsquo;wing length\u0026rsquo;, and \u0026lsquo;body mass\u0026rsquo;, as well as a random intercept for the grouping variable \u0026lsquo;sex\u0026rsquo;.\u003c/p\u003e \u003cp\u003eIn the Global Messenger software, we exported the satellite-tracked migration route data in .shp format. Subsequently, we added the downloaded imagery layer from Google Earth along with the migration route shp layer to ArcGIS 10.6 to delineate the spring and autumn migration routes of the Common cuckoo. Migration routes of different individuals are depicted in distinct colors, breeding sites are represented by rectangles, and wintering sites are denoted by triangles. Stopover sites are categorized based on the duration of stay, ranging from 2\u0026ndash;7 days, 7\u0026ndash;14 days, 14\u0026ndash;21 days, to more than 21 days, with different-sized circles representing different types of mid-stopover sites. We conducted independent sample t-tests to compare migration parameters among different routes, genders, and seasons. Data for each group are presented as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error. The data processing was carried out using R version 4.2.2.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eMigration routes\u003c/h2\u003e \u003cp\u003eWithin the study region, migration routes of 13 Common cuckoos (comprising 7 males and 6 females) were systematically documented from 2018 to 2021 (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Notably, two individuals, identified as 201907 and 202115, achieved successful completion of the autumnal migratory endeavor. Additionally, a singular representative, denoted as 201907, undertook a successful spring migration. Population of common cuckoo breeding in Central Asia manifest three principal autumn migration routes (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), Specifically, 8 individuals (61.5% of the total) opted for the western route around the Qinghai-Tibet Plateau and the Taklamakan Desert. Notably, 201907 undertakes two consecutive autumn migrations, ultimately wintering in Tanzania. 2 Common cuckoos (15.4% of the total) choosed to traverse the Qilian Mountains from their breeding sites, proceeding southward over the Qinghai-Tibet Plateau via Xining. However, 1 individual lost its signal near Lhasa after reaching that vicinity. 3 common cuckoos (23.1%) selected for the eastern route, circumventing the eastern part of the Qinghai-Tibet Plateau. Notably, Common cuckoo 202115 successfully completed its autumn migration and accomplished wintering in Mozambique (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The difference in wintering locations between these two common cuckoos (i.e. identified as 201907, 202115) from the same breeding site implies, to some extent, a relatively weak migratory connectivity within the common cuckoo population.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eCommon cuckoo Individuals Tracked from 2018 to 2021\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"10\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSeason\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBirds ID\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTracker ID\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGender\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eDeparture date\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eArrival date\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eMigration distance (Km)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eTarsus(mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eWing(mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eWeight(g)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"13\" rowspan=\"14\"\u003e \u003cp\u003eAutumn\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e201801\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eB008\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♂\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2018.8.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2268.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e22.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e180\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e98\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e201803\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eB006\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♀\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2018.8.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2045.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e25.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e215\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e104.08\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e201804\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eB007\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♀\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2018.8.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e3279.48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e23.48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e200\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e92.86\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e201805\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eB010\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♂\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2018.7.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e1125.68\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e23.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e219\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e95.91\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e201906\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eB011\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♀\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2019.7.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e6248.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e23.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e205\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e90.67\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e201907(1st)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eB012\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♀\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2019.7.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2019.11.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e9459.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e24.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e201.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e95.26\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e201907(2nd)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eB012\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♀\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2020.7.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2020.11.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e9840.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e202010\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eB027\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♀\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2020.7.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e1288.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e21.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e197\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e92.95\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e202111\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHQP3660\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♂\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2021.7.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e4707.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e23.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e212\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e110.04\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e202113\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHQP3662\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♂\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2021.7.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2280.85\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e22.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e216\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e101.8\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e202114\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHQP3663\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♀\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2021.7.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2618.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e25.73\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e210\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e126.59\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e202115\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHQP3667\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♂\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2021.8.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2021.12.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e11594.56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e23.61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e211\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e101.7\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e202116\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHQP3669\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♂\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2021.8.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e6748.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e23.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e207\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e111.78\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e202117\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHQP3671\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♂\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2021.8.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2139.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e24.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e212\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e107.16\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSpring\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e201907\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eB012\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♀\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2020.3.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2020.5.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e9033.51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e202115\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHQP3667\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e♂\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2022.4.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e9197.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eIn the western route of autumn migration, the most utilized stopover site is situated in Hotan, Xinjiang (80.78E, 36.25N), with four Common cuckoos (identified as 201801, 201907, 202116, 202117) choosing this site for resting. On the eastern autumn migration route, the secondary hub for stopovers is Baoding City, Yunnan, China (99.23E, 25.46N), where three Common cuckoos (identified as 202010, 202103, 202115) were observed resting. In the central migration route, both Common cuckoos paused for rest in Delingha City, Qinghai, China (98.26E, 36.62N). Furthermore, Dinder National Park in Sudan (34.91E, 12.23N) represens the stopover site with the longest duration (46.5 days). Following the completion of spring migration, Common cuckoo 201907 did not return to its original breeding site in Guazhou County. Instead, it selected a new breeding site near Barkol Kazakh Autonomous County, Hami City, Xinjiang (91.19\u0026ndash;94.48 E, 43.21\u0026ndash;45.05N) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). During its second autumn migration, this individual chose stopover sites in Pakistan (Mangla, 73.59E, 33.17N), Eritrea (Gash-Barka, 37.56E, 15.27N), and Sudan (Dinder National Park, 34.91E, 12.23N), all of which are characterized by latitudinal proximity (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe main stopover sites and durations during the migration process of the Central Asian breeding population of Common cuckoos.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"9\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSeason\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eStopover site\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eRegion\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eLongitude\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLatitude\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eBirds\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eArrive time\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eDeparture time\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eDuration (day)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAutumn\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHotan Prefecture\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eChina\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e80.78E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e36.25N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201801\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2018.8.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2018.8.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e5.37\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201907\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2019.8.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2019.8.10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e5.75\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e202116\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2021.8.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2021.8.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e12.17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e202117\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2021.8.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2021.8.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e8.25\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eKashi area\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eChina\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e76.49E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e38.66N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201804\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2018.9.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2018.9.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e15.12\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDelingha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eChina\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e98.26E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e36.62N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201803\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2018.8.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2018.8.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e11.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201805\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2018.8.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2018.8.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e10.25\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBaoshan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eChina\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e99.23E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e25.46N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e202010\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2020.8.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e20208.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e18.25\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e202113\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2021.8.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2021.8.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e20.75\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e202115\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2021.8.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2021.9.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e21.75\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMangla\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePakistan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e73.59E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e33.17N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201907(1st)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2019.8.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2019.8.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e14.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201907(2nd)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2020.7.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2020.8.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e24.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e202116\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2021.8.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2021.9.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e13.42\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePathankot\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIndia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e75.81E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e35.54N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201906\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2019.8.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2019.9.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e22\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGash-Barka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEritrea\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e37.56E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e15.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201907(1st)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2019.9.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2019.9.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e14.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201907(2nd)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2020.9.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2019.9.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e12.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDinder National Park\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSudan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e34.91E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e12.23N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201907(1st)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2019.9.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2019.11.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e46.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201907(2nd)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2020.10.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2020.11.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e26.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSpring\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMambasa\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCongo\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e29.70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201907\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2020.4.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2020.4.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e13.25\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAgew Awi Zone\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEthiopia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e36.29E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e10.56N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201907\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2020.4.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2020.4.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e7.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarayan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIran\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e58.19E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e33.18N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e201907\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2020.5.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2020.5.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e6.25\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLuuq\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSomalia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e42.77E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3,34N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e202115\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2022.4.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2021.4.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e4.25\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eMigrations traits\u003c/b\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe results indicated that the Autumn migration distance of 201907 for the first and second subjects were 9459.45km and 9840.14km, respectively. However, the Autumn migration distance of 202115 was 11594.56km. We used independent sample t-test to compare Common cuckoo's autumn and spring migration (Fig.\u0026nbsp;3), a significant difference emerged in the migratory speed. Specifically, the spring migration speed was notably higher than that observed in autumn migration (spring speed: 789.80\u0026thinsp;\u0026plusmn;\u0026thinsp;314.68 km/day, autumn speed: 548.61\u0026thinsp;\u0026plusmn;\u0026thinsp;340.37 km/day, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.002). Furthermore, the duration of stopover at intermediate sites during spring migration was significantly shorter compared to autumn migration (spring stopover duration: 9.97\u0026thinsp;\u0026plusmn;\u0026thinsp;7.16 days, autumn stopover duration 13.28\u0026thinsp;\u0026plusmn;\u0026thinsp;8.11 days, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.01). However, there was no statistically significant difference in migratory straightness between spring and autumn migrations (spring straightness: 0.88\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13, autumn straightness: 0.88\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.21).\u003c/p\u003e \u003cp\u003eStatistical analysis of instantaneous speed data from 13 common cuckoos during migration in the period of 2018\u0026ndash;2021, excluding data points with instantaneous speeds below 10 km/h, revealed distinct patterns. During autumn migration (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e4\u003c/span\u003e), Common cuckoo migration predominantly occurred during the night, concentrated from 0:00 to 6:00 (84.77% of the total migration time). In contrast, during spring migration, the daily migration time span was concentrated from 0:00 to 12:00 (89.08%). There was no significant variation in the selection of migration routes based on the tarsus length and body mass of common cuckoos. However, wing length have a significant difference between various migration routes. Wing length of Common cuckoos that select middle and east migration routes is longer than individuals of west migration routes (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe impact of migration route selection on the response of morphological traits.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colspan=\"7\" nameend=\"c7\" namest=\"c1\"\u003e \u003cp\u003elinear mixed model parameters\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFixed effects\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eβ\u0026thinsp;\u0026plusmn;\u0026thinsp;SE\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003edf\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003et\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e\u003cem\u003eP\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eIntercept\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.893\u0026thinsp;\u0026plusmn;\u0026thinsp;3.115\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-0.287\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e0.781\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTarsus length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.181\u0026thinsp;\u0026plusmn;\u0026thinsp;0.149\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-1.216\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e0.255\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWing length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.033\u0026thinsp;\u0026plusmn;\u0026thinsp;0.014\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.311\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e0.046\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBody mass\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.012\u0026thinsp;\u0026plusmn;\u0026thinsp;0.016\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-0.765\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e0.464\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRandom effects\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003eβ\u0026thinsp;\u0026plusmn;\u0026thinsp;SD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003en\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eResults of VCA (%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003esex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e0.001\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eResidual\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e0.199\u0026thinsp;\u0026plusmn;\u0026thinsp;0.447\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e99.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eSE of fixed effects is the standard error of the mean; SD of random effects is the square root of the variance.\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eOur findings reveal three primary autumn migration routes in the Central Asian common cuckoo population: the western route initiates along the southern edge of the Taklamakan Desert and the western edge of the Himalayas, the central route traverses the Tibetan Plateau, and the eastern route follows the edge of the Tibetan Plateau. The variation of migration routes suggests a significant impact of the Qinghai-Tibet Plateau, a large ecological barrier, on the common cuckoo, leading to a high variation in their migratory routes. Different species populations may adopt similar strategies when facing same large ecological barriers like the Tibetan Plateau. Juhant and Bildstein (2017) [36] identified four main migration corridors: the western Himalayan corridor, the eastern Himalayan corridor, the north-south corridor, and the corridor traversing the Himalayas in their study on raptor migration through the Tibetan Plateau. Correspondingly, Migration routes of black kites are similar with the west route [37]. These routes encompass traversing the Taklamakan Desert and the Tianshan Mountains before arriving at their breeding sites. Regarding the middle routes of autumn migration is not fully realized, there is a discernible inclination toward crossing, and the potential for common cuckoo to detour to the southern part of the Tibetan Plateau later in the season is not excluded. Lee et al. [38] discovered that the cuckoo population breeding in Korea migrates to Southeast Africa during the winter, exhibiting migration routes and wintering areas like those of the eastern route of the Central Asian population. Conversely, common cuckoos breeding in Beijing follow a route around the southern edge of the Qinghai-Tibetan Plateau, resembling the western route in autumn migration. Consequently, we posited various species or populations, when confronted with the same ecological barrier, exhibit a remarkable convergence in their selection of migratory routes. La Sorte and Fink [39] based on information from the eBird platform, analyzed seven years of migration routes for 55 terrestrial bird species. They found that ecological barriers increase the flexibility of migratory routes for birds, as exemplified by the diverse choices of migration routes among these 55 species across the American continent.\u003c/p\u003e \u003cp\u003eIndividuals of Common cuckoo on the same migration route choose identical stopover sites which may be associated with environmental factors during the migration process. Thorup et al. [40] observed a propensity for Common cuckoos to select regions characterized by elevated vegetation cover during migration in their investigation of migratory bird dynamics and resource tracking. Another pivotal determinant could be the shared encounter of individuals on the same migratory route with homogeneous ecological impediments. The presence of substantial ecological barriers has the potential to protract the migratory duration for migrant birds engaged in long-distance flights with migration timings and routes demonstrating seasonal differentiations. The existence of such ecological barriers serves to prompt migratory birds to converge upon designated intermediate rest sites for strategic energy storage and replenishment [39]. During its second autumn migration, the selection of the same stopover sites by 201907 was postulated to be linked to its protracted memory retention. Migrant birds engaged in extensive long-distance flights often exhibit an extended duration of memory retention for stopover sites [41\u0026ndash;43]. Research in the peregrine populations of the Eurasian Arctic revealed that long-term memory emerged as the most probable selective agent for the divergence observed in ADCY8 [44].\u003c/p\u003e \u003cp\u003eBetween different seasons, the speed of spring migration is significantly higher than that of autumn migration, and the duration of stopovers is notably shorter in spring than in autumn. Analyzing data with instantaneous speeds exceeding 10 km/h, we also observed that the common cuckoo exhibits typical nocturnal migratory behavior, with longer daily flight times during spring migration compared to autumn migration. Birds generally prefer migrating with tailwinds [45], as seen in species such as the European Honey Buzzard (\u003cem\u003ePernis apivorus\u003c/em\u003e), White Stork (\u003cem\u003eCiconia ciconia\u003c/em\u003e), and Western Osprey (\u003cem\u003ePandion haliaetus\u003c/em\u003e). The strength of tailwinds is higher in spring than in autumn, providing a plausible explanation for the observed phenomenon of migratory birds exhibiting faster migration speeds during the spring compared to the autumn season [46]. The faster migratory speed and reduced stopover duration observed in the common cuckoo during spring migration suggested the adoption of a time-minimization strategy. This spring migration strategy aligns with that commonly observed in many avian species, wherein the pursuit of a time-minimization approach allows birds to secure high-quality breeding habitats earlier, thus enhancing reproductive success [47, 48]. Additionally, this strategy diminishes the frequency and duration of stopovers during migration, thereby mitigating the risk of predation by avian predators [49]. Given the common cuckoo's status as a brood parasite without nest-building behaviors, the time minimization strategy employed during spring migration may be linked to the migratory tactics of its hosts. Birds of the Shrike family (\u003cem\u003eLaniidae\u003c/em\u003e) exhibited faster spring migration speeds compared to autumn migration, and their selection of stopover sites tends to favor areas with abundant vegetation. During spring migration, these birds adopt a strategy that minimizes time spent in transit, aiming to reach their breeding grounds quickly and aligning with the peak of vegetation coverage upon arrival [50\u0026ndash;52]. Research has indicated common cuckoos tend to initiate its migration approximately a month later than its parasitic host [53]. Therefore, the expedited arrival at breeding grounds during spring migration in common cuckoos may be construed as an adaptive response aligning with the reproductive timing of their hosts.\u003c/p\u003e \u003cp\u003eOur results indicate that individuals opting for eastward and middle migration routes exhibit longer wing length compared to those selecting westward routes. Since the eastern and middle routes are longer than the western routes, Individuals choosing the eastern and central routes may possess longer wings to improve flight efficiency, thus facilitating their ability to cover longer migration distances. This difference has been observed in other migrant birds, Eurasian Blackcap (\u003cem\u003eSylvia atricapilla\u003c/em\u003e) has been reported to exhibit various migration routes among individuals, a phenomenon attributed to differences in wing length [54]. Hahn et al. [55] found that longer wing provides benefits in adjusting the speed of migrants to their flight and arrival time. Additionally, the habitats along western migration routes mainly consist of deserts and high mountains, while eastern migration routes offer relatively abundant habitats. Therefore, this result suggests a potential trade-off between morphological traits and the availability of better stopover sites. The reason why the eastern and middle migration routes, which involve longer migration distances, are chosen by cuckoos with relatively longer wing length may be that longer wings provide them with higher flight efficiency and better selection of stopover sites along the way.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eOur study revealed that the migration routes of the common cuckoos breeding in Central Asia are influenced by the Qinghai-Tibet Plateau. It demonstrated the migration strategy of the common cuckoo and suggested that their selection of migration routes may be influenced by wing length. Constrained by data limitations, the investigation into the migratory behavior of the common cuckoo population breeding in Central Asia necessitates ongoing efforts. Further substantiation is required for tracking the middle routes during autumn migration.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAvailability of data and materials\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets analysed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank the management staff of Gansu An\u0026rsquo;xi Extremely arid Desert National Nature Reserve for allowing and helping us to conduct this study in the area. We also appreciate the support from Hunan Global Messenger company.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFinancial support of this study was provided by the National Natural Science Foundation of China (Grants 31672296).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe study was approved by the Ethics Committee of School of Life Sciences of Lanzhou University (File no 20180330), and strictly complied with the requirements of the Chinese Wild Animal Protection Law.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no conflicts of interest to any other affiliations or persons.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eContributions:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eZC wrote this manuscript, developed most of the data summary, and conducted the data analysis. DM and LK participated in data analysis and field activities. ML and XD collected the filed data. XB designed the experiments and refined this manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n \u003cli\u003e\u003cspan\u003eAlerstam T, Hedenstr\u0026ouml;m A, \u0026Aring;kesson S: \u003cstrong\u003eLong-distance migration: evolution and determinants\u003c/strong\u003e. \u003cem\u003eOikos\u003c/em\u003e 2003, \u003cstrong\u003e103\u003c/strong\u003e(2):247\u0026ndash;260.\u003c/span\u003e\u003c/li\u003e\n \u003cli\u003e\u003cspan\u003eLack D: \u003cstrong\u003eBird migration and natural selection\u003c/strong\u003e. \u003cem\u003eOikos\u003c/em\u003e 1968:1\u0026ndash;9.\u003c/span\u003e\u003c/li\u003e\n \u003cli\u003e\u003cspan\u003eWebster MS, Marra PP, Haig SM, Bensch S, Holmes RT: \u003cstrong\u003eLinks between worlds: unraveling migratory connectivity\u003c/strong\u003e. \u003cem\u003eTrends in ecology \u0026amp; evolution\u003c/em\u003e 2002, \u003cstrong\u003e17\u003c/strong\u003e(2):76\u0026ndash;83.\u003c/span\u003e\u003c/li\u003e\n \u003cli\u003e\u003cspan\u003eHerrera CM: \u003cstrong\u003eOn the breeding distribution pattern of European migrant birds: MacArthur\u0026apos;s theme reexamined\u003c/strong\u003e. \u003cem\u003eThe Auk\u003c/em\u003e 1978, \u003cstrong\u003e95\u003c/strong\u003e(3):496\u0026ndash;509.\u003c/span\u003e\u003c/li\u003e\n \u003cli\u003e\u003cspan\u003eKubelka V, \u0026Scaron;\u0026aacute;lek M, Tomkovich P, V\u0026eacute;gv\u0026aacute;ri Z, Freckleton RP, Sz\u0026eacute;kely T: \u003cstrong\u003eGlobal pattern of nest predation is disrupted by climate change in shorebirds\u003c/strong\u003e. \u003cem\u003eScience\u003c/em\u003e 2018, \u003cstrong\u003e362\u003c/strong\u003e(6415):680\u0026ndash;683.\u003c/span\u003e\u003c/li\u003e\n \u003cli\u003e\u003cspan\u003eNewton I, Dale L: \u003cstrong\u003eRelationship between migration and latitude among west European birds\u003c/strong\u003e. \u003cem\u003eJournal of Animal Ecology\u003c/em\u003e 1996:137\u0026ndash;146.\u003c/span\u003e\u003c/li\u003e\n \u003cli\u003e\u003cspan\u003eNewton I, Dale L: \u003cstrong\u003eBird migration at different latitudes in eastern North America\u003c/strong\u003e. \u003cem\u003eThe Auk\u003c/em\u003e 1996, \u003cstrong\u003e113\u003c/strong\u003e(3):626\u0026ndash;635.\u003c/span\u003e\u003c/li\u003e\n \u003cli\u003e\u003cspan\u003eBoth C, Van Turnhout CA, Bijlsma RG, Siepel H, Van Strien AJ, Foppen RP: \u003cstrong\u003eAvian population consequences of climate change are most severe for long-distance migrants in seasonal habitats\u003c/strong\u003e. \u003cem\u003eProceedings of the Royal Society B: Biological Sciences\u003c/em\u003e 2010, \u003cstrong\u003e277\u003c/strong\u003e(1685):1259\u0026ndash;1266.\u003c/span\u003e\u003c/li\u003e\n \u003cli\u003e\u003cspan\u003eSherry TW, Wilson S, Hunter S, Holmes RT: \u003cstrong\u003eImpacts of nest predators and weather on reproductive success and population limitation in a long-distance migratory songbird\u003c/strong\u003e. \u003cem\u003eJournal of avian biology\u003c/em\u003e 2015, \u003cstrong\u003e46\u003c/strong\u003e(6):559\u0026ndash;569.\u003c/span\u003e\u003c/li\u003e\n \u003cli\u003e\u003cspan\u003eLok T, Overdijk O, Piersma T: \u003cstrong\u003eThe cost of migration: spoonbills suffer higher mortality during trans-Saharan spring migrations only\u003c/strong\u003e. \u003cem\u003eBiology letters\u003c/em\u003e 2015, \u003cstrong\u003e11\u003c/strong\u003e(1):20140944.\u003c/span\u003e\u003c/li\u003e\n \u003cli\u003e\u003cspan\u003eDeppe JL, Ward MP, Bolus RT, Diehl RH, Celis-Murillo A, Zenzal Jr TJ, Moore FR, Benson TJ, Smolinsky JA, Schofield LN: \u003cstrong\u003eFat, weather, and date affect migratory songbirds\u0026rsquo; 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\u003cstrong\u003e6\u003c/strong\u003e(1):68\u0026ndash;77.\u003c/span\u003e\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","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":"Cuculus canorus, birds migration, Qinghai-Tibet Plateau, migration routes, migrant strategy","lastPublishedDoi":"10.21203/rs.3.rs-3967127/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3967127/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eSeasonal climate changes and fluctuations in food resources drive the migratory behavior of birds between their breeding and wintering sites. Migratory birds change their migration routes in response to climate and environmental stresses. Existing studies have indicated that the widely distributed Eurasian Common cuckoo migrates individually during the night, necessitating further research to elucidate its global migration routes. From 2018 to 2021, the migration routes of 13 adult Common cuckoos breeding in the north of the Qinghai-Tibet Plateau were tracked using satellite trackers. We found that (1) the migration routes of this Common cuckoo population mainly separate into three directions: 8 individuals followed the western routes along the western edge of the Qinghai-Tibet Plateau to the south, 3 individuals traversed the eastern routes along the eastern part of the Qinghai-Tibet Plateau and the rest 2 opted for the middle routes across most of the Qinghai-Tibet Plateau to the south. (2) Individual marked with the identifier 201907 exhibited a consistent stopover site selection during autumn migration. However, it changed the breeding area after spring migration. (3) In the context of autumn migration, 4 tracked Common cuckoos (50%) along the western migratory routes exhibited a consistent preference for the Hotan area in Xinjiang as their stopover site. Concurrently, 3 individuals (100%) traversing the eastern migratory route consistently selected stopover locations in the vicinity of Baoshan city, Yunnan. (4) The migration speed of Common cuckoos is significantly higher in spring than in autumn, and the stopover duration of spring migration is significantly lower than that of autumn migration. The daily flight time of spring migration is concentrated between 0\u0026ndash;12 clock, while that of autumn migration is between 0\u0026ndash;6 clock. (5) Migration routes of the Central Asian Common cuckoo population were limited by the Qinghai-Tibet plateau. The population exhibited high levels of plasticity, with individuals with longer wings tending to select the east and middle routes, while shorter wings were observed more frequently in the west route. Our findings reveal that central Asian common cuckoo populations exhibit a preference for a minimum-time strategy during spring migration, with migration routes selected based on wing length.\u003c/p\u003e","manuscriptTitle":"Variation of migration routes in the Central Asian-breeding Cuculus canorus population influenced by the Qinghai-Tibet Plateau","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-02-26 20:24:09","doi":"10.21203/rs.3.rs-3967127/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":"82a28061-98b5-4c23-8e70-add6ac715c4c","owner":[],"postedDate":"February 26th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2024-03-01T17:37:23+00:00","versionOfRecord":[],"versionCreatedAt":"2024-02-26 20:24:09","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-3967127","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3967127","identity":"rs-3967127","version":["v1"]},"buildId":"FbvkV6FR0MCFSLy54lSbu","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}