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Cuculus Canorus): Isolation by Distance and Divergence Among Subspecies

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Cuculus Canorus): Isolation by Distance and Divergence Among Subspecies J Ornithol (2015) 156:533–542 DOI 10.1007/s10336-014-1153-6 ORIGINAL ARTICLE Geographic variation in the calls of the Common Cuckoo (Cuculus canorus): isolation by distance and divergence among subspecies Chentao Wei • Chenxi Jia • Lu Dong • Daiping Wang • Canwei Xia • Yanyun Zhang • Wei Liang Received: 3 June 2014 / Revised: 8 November 2014 / Accepted: 19 December 2014 / Published online: 13 January 2015 Ó Dt. Ornithologen-Gesellschaft e.V. 2015 Abstract Studies on the pattern of geographic variation distance and environmental differences on call differenti- in bird vocalizations can facilitate the understanding of the ation. The results showed there to be significant differences evolutionary history of species and species differentiation. in the calls of different subspecies of the Common Cuckoo. The Common Cuckoo (Cuculus canorus) is a non-passer- Discriminant function analysis was able to correctly iden- ine widely distributed in Eurasia, and its calls are not tify 81.7 % of individuals to their original subspecies, and acquired through learning. Revealing the geographic pat- 98 % of individuals of subspecies subtelephonus were tern of Common Cuckoo calls may help our understanding correctly assigned. Differences in calls both within and of the relationship between environment, genetic differ- between subspecies were found to be significantly corre- entiation, and vocal differentiation. In the present study, lated with geographic distance, while environmental dif- geographic variation in the calls of the Common Cuckoo ferences have no important effect. Our study stressed the was investigated throughout Eurasia for the first time. Calls effect of isolation by distance on geographic variation of of different subspecies of the Common Cuckoo were non-passerine vocalization, and we infer that the great compared, and the correlations between differences in divergence in calls between different Common Cuckoo calls, geographic distance, climatic differences, and alti- subspecies may be a hint of cryptic species. tude differences were determined in order to evaluate the influence of subspecies differentiation, isolation by Keywords Common Cuckoo Á Cuculus canorus Á Eurasia Á Call variation Á Subspecies Á Isolation by distance Communicated by F. Bairlein. Zusammenfassung Electronic supplementary material The online version of this article (doi:10.1007/s10336-014-1153-6) contains supplementary Geographische Variation in den Rufen des Kuckucks material, which is available to authorized users. (Cuculus canorus): Isolation durch Entfernung und C. Wei Á L. Dong Á D. Wang Á C. Xia Á Y. Zhang Unterschiede zwischen Unterarten Ministry of Education Key Laboratory for Biodiversity Science and Ecological Engineering, College of Life Sciences, Studien u¨ber die geographischen Verteilungsmuster von College of Life Sciences, Beijing Normal University, 100875 Beijing, China Lauta¨ußerungen der Vo¨gel ko¨nnen das Versta¨ndnis der evolutiona¨ren Geschichte von Arten erleichtern. Der Ku- C. Wei Á W. Liang (&) ckuck (Cuculus canorus) ist ein in Eurasien weit verbre- Ministry of Education Key Laboratory for Tropical Animal and iteter Nichtsperlingsvogel, dessen Rufe nicht erlernt Plant Ecology, College of Life Sciences, Hainan Normal University, 571158 Haikou, China werden. Die geographische Verteilung der Kuckucksrufe e-mail: [email protected] darzustellen ko¨nnte helfen den Zusammenhang zwischen Umwelt, genetischer Differenzierung und stimmlicher C. Jia Differenzierung zu verdeutlichen. In dieser Studie wurden Key Laboratory of the Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, erstmals die geographischen Unterschiede zwischen den 100101 Beijing, China Rufen des Kuckucks in Eurasien untersucht. Die Rufe 123 534 J Ornithol (2015) 156:533–542 verschiedener Unterarten wurden verglichen und die Kor- Vocal divergence also occurs in populations that expe- relationen zwischen Unterschieden im Ruf, geographischer rience prolonged isolation (Koetz et al. 2007; Robin et al. Entfernung, klimatischer Unterschiede und Ho¨henunters- 2011; Parker et al. 2012). Genetic and cultural drift can chieden berechnet, um die Einflu¨sse von Unterartendiffer- lead to vocal differentiation, and geographic isolation can enzierung, Isolation durch Entfernung und Umweltfaktoren facilitate the accumulation of these differences in different auf Rufe zu ermitteln. Es zeigten sich signifikante Unter- populations. Subspecies differentiation represents histori- schiede in den Rufen verschiedener Unterarten des Ku- cal isolation between populations, therefore relatively large ckucks. Eine Diskriminanzanalyse war in der Lage, 81,7 % vocal differences are also prone to occur (Dingle et al. der Individuen ihrer Unterart richtig zuzuordnen und 98 % 2008; Turcokova et al. 2010). der Individuen der Unterart subtelephonus wurden richtig The Common Cuckoo (Cuculus canorus) is a migratory zugeordnet. Unterschiede in Rufen sowohl in derselben brood parasite. It breeds extensively in Eurasia and a few Unterart als auch zwischen Unterarten waren signifikant places in North Africa and winters in Africa and southern mit der geographischen Entfernung korreliert, wa¨hrend Asia. There are four Common Cuckoo subspecies, canorus, Umweltfaktoren keinen Einfluss hatten. Unsere Studie subtelphonus, bakeri, and bangsi. They differ notably in betont den Effekt der Isolation durch Entfernung auf die plumage color (Payne 2005). The Common Cuckoo is a geographischen Unterschiede in den Lauta¨ußerungen von non-passerine bird and during the breeding season, the Nichtsperlingsvo¨geln und wir schließen aus der großen males emit loud, characteristic ‘‘cu-coo’’ calls. It is gen- Variation in Rufen zwischen verschiedenen Unterarten des erally considered that its call does not vary very much Kuckucks, dass es sich um eine kryptische Art handeln across the entire distribution (Payne 2005). However, ko¨nnte. studies on the variation of Common Cuckoo calls have been either on a small-scale (Fuisz and de Kort 2007; Jung et al. 2014) or limited by a small sample size (Lei et al. 2005). There has been no investigation on the pattern of Introduction geographic variation in the calls of the Common Cuckoo over a large geographic area, and none of the previous Vocal cues play an important role in species recognition and studies discussed vocal divergence between subspecies. mate choice in birds (Marler and Slabbekoorn 2004; Catch- This species is widely distributed in Eurasia, where envi- pole and Slater 2008). Differences in vocalization among ronments can differ greatly. In addition, this cuckoo may different geographic populations can sometimes lead to be the only non-passerine species with a distribution which reproductive isolation, thereby promoting the formation of extends from sea level (in Europe and on the east coast of new species (Irwin 2000). Consequently, studies on geo- China) to high altitudes (up to 4,000 m) in the Himalayas graphic variation in songs and calls of birds are of significant and southwest China. The acoustic adaptation hypothesis importance in understanding the relationship between mating predicts that call properties are adapted to different envi- signal evolution and species differentiation. ronments. If so, Common Cuckoo calls may show rela- The song divergence of many songbirds can be tively large differences between environments and may described using the isolation by distance model (Irwin have a close correlation with environmental variables. 2000; Rivera-Gutierrez et al. 2010; Xing et al. 2013). It is According to the isolation by distance model, large dis- generally believed that these patterns are caused by cul- tances reduce the likelihood of gene flow between popu- tural drift and that differences in songs increase with lations (Wright 1943). Because the Common Cuckoo is a distance due to inaccuracies in the learning process. non-passerine, its call is affected more by genetic factors However, studies examining whether the differentiation in than by other factors such as the learning process, while the calls of suboscine and non-passerine fit the isolation genetic differentiation caused by isolation by distance may by distance model have been rare (Pe´rez-Mena and Mora also lead to vocal divergence between remote places. 2011; Lovell and Lein 2013). Non-learned calls may also Meanwhile, subspecies differentiation represents a past fit the isolation by distance model. For example, a study history of vicariance events, and differences in plumage on the calls of the Greenish Warbler (Phylloscopus tro- color indicate relatively large genetic differences between chiloides) showed that the call differentiation is correlated different Common Cuckoo subspecies, so it is also quite with geographic and genetic differences (Irwin et al. possible that there will be large vocal divergences among 2008). The calls of non-passerines are not acquired them. It has been suggested that, within the Common through learning and are therefore not subject to cultural Cuckoo species, there may exist several genetically drift. Whether the differentiation of their calls can still be divergent gentes (Gibbs et al. 2000; Fossøy et al. 2011). A described by the isolation by distance model is worth study showed there to be significant differences in the calls exploring. of the Common Cuckoo in different habitats where they 123 J Ornithol (2015) 156:533–542 535 parasitise different hosts (Fuisz and de Kort 2007). To fully Collection of climate and altitude data understand the geographic
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