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Mongolian Short-Toed Lark Calandrella Dukhunensis, An

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Mongolian Short-Toed Lark Calandrella Dukhunensis, An Journal of Ornithology (2021) 162:165–177 https://doi.org/10.1007/s10336-020-01819-z ORIGINAL ARTICLE Mongolian Short‑toed Lark Calandrella dukhunensis, an overlooked East Asian species Per Alström1,2 · Gombobaatar Sundev3 Received: 6 May 2020 / Revised: 27 August 2020 / Accepted: 8 September 2020 / Published online: 23 September 2020 © The Author(s) 2020 Abstract The eastern subspecies of Greater Short-toed Lark Calandrella brachydactyla dukhunensis has recently been considered a separate species, Calandrella dukhunensis, by several authors based on molecular data. We present supporting evidence for this treatment based on studies of morphology, vocalisations and song-fight, and also present new data on other aspects of its biology based on feld studies. We show that its breeding distribution is considerably smaller than previously thought, and is restricted to the eastern half of Mongolia and, marginally, neighbouring parts of China and perhaps Russia. Keywords Calandrella brachydactyla · Taxonomy Zusammenfassung Die Chinalerche Calandrella dukhunensis, eine übersehene ostasiatische Art Die östliche Unterart der Kurzzehenlerche Calandrella brachydactyla dukhunensis wird seit kurzem von mehreren Autoren auf Basis molekularer Daten als eigene Art, Calandrella dukhunensis, betrachtet. Wir legen hier zusätzliche Belege für diese Einstufung vor. Diese beruhen auf Untersuchungen zur Morphologie, Vokalisation und des Singfuges. Basierend auf Feldstudien präsentieren wir auch neue Daten über andere Aspekte der Biologie dieser Art. Wir zeigen, dass das Brutgebiet wesentlich kleiner ist, als bisher angenommen und sich auf die östliche Hälfte der Mongolei und kleine Teile des benachbarten Chinas und eventuell Russlands beschränkt. Introduction The Greater Short-toed Lark Calandrella brachydactyla was previously considered conspecifc with Red-capped Lark C. cinerea, under that name (e.g. Meinertzhagen 1951; Vaurie Communicated by J. T. Lifeld. 1959; Peters 1960), but was later suggested to be specif- cally diferent (reviewed in Stervander et al. 2016). Caland- Electronic supplementary material The online version of this rella brachydactyla was considered to breed from northwest article (https ://doi.org/10.1007/s1033 6-020-01819 -z) contains supplementary material, which is available to authorized users. Africa and the Iberian Peninsula through southern Europe and western and central Asia to northeastern Mongolia and * Per Alström adjacent parts of Russia and China, and across the eastern [email protected] part of the Tibetan Plateau to just north of the Himalayas, 1 Animal Ecology, Department of Ecology and Genetics, and to winter locally in northwestern Africa, but mainly in Evolutionary Biology Centre, Uppsala University, the Sahel region of Africa and the Arabian Peninsula east to Norbyvägen 18D, 752 36 Uppsala, Sweden India (Glutz von Blotzheim and Bauer 1985; Cramp 1988; 2 Key Laboratory of Zoological Systematics and Evolution, Sibley and Monroe 1990; de Juana and Suárez 2004; Dick- Institute of Zoology, Chinese Academy of Sciences, inson and Christidis 2014; Christidis 2018). Most authors Beijing 100101, China recognised 6–9 subspecies. 3 National University of Mongolia and Mongolian Alström et al. (2013) found, based on the mitochon- Ornithological Society, Ulaanbaatar 210646a, P.O. Box 537, drial cytochrome b gene, that the eastern subspecies, C. b. Ulaanbaatar, Mongolia Vol.:(0123456789)1 3 166 Journal of Ornithology (2021) 162:165–177 dukhunensis, was more closely related to Hume’s Lark C. song-fight, breeding, and migration period of this species acutirostris than to other populations of C. brachydactyla based on feld work and studies of museum specimens. (as well as C. cinerea). This was corroborated by Stervander et al. (2016) based on genomic data for the same single C. b. dukhunensis sample analysed by Alström et al. (2013), Materials and methods collected on the breeding grounds in eastern Mongolia, as well as cytochrome b sequence data from fve additional C. Throughout, we treat Calandrella dukhunensis as a sepa- b. dukhunensis collected in the winter quarters in central rate species from C. brachydactyla, based on the phyloge- and northeast India. The divergence time between western netic analyses by Alström et al. (2013) and Stervander et al. taxa of C. brachydactyla and C. b. dukhunensis plus C. (2016). For subspecies taxonomy of C. brachydactyla, we acutirostris was estimated at 6.0 million years ago (mya) follow del Hoyo et al. (2020). G.S. has studied C. dukhun- by Alström et al. (2013) and c. 5 mya by Stervander et al. ensis in Mongolia on multiple occasions during 2005–2019, (2016), although their confdence intervals for the dates were together with P.A. in mid-June 2016. P.A. has also observed broadly overlapping. this species multiple times on spring and autumn migra- The results from the phylogenetic analyses have led some tion in China, and has examined specimens in the American recent authors to recognise C. dukhunensis as a distinct spe- Museum of Natural History, New York, USA (AMNH); Uni- cies, with the English name Sykes’s Short-toed Lark (Cle- versity of Michigan Museum of Zoology, Ann Arbor, USA ments et al. 2019), Mongolian Short-toed Lark (Shirihai and (UMMZ); Natural History Museum, Tring, UK (NHM); Svensson 2018; Gill et al. 2020) or Eastern Short-toed Lark National Zoological Museum of China, Institute of Zool- (del Hoyo and Collar 2016; del Hoyo et al. 2020). How- ogy, Chinese Academy of Sciences, Beijing (NZMC-IOZ); ever, there is no consensus among these authors regarding and Yamashina Institute of Ornithology, Chiba, Japan (YIO; its breeding range. According to Clements et al. (2019), it on loan). In addition, P.A. has studied C. brachydactyla at breeds in “north central China (central and eastern Xinjiang) multiple places throughout its breeding distribution (e.g. south to Tibet”; according to Shirihai and Svensson (2018) Morocco, Kazakhstan, northwest China) and on migration “Tibetan plateau, and through E Xinjiang and S and E Mon- and in winter at multiple places (e.g. northwest India), as golia to E Transbaikalia and W Manchuria”; according to well as in the above and other museum collections (e.g. Gill et al. (2020) in west and central China; and according University of Copenhagen Zoological Museum, Denmark; to del Hoyo and Collar (2016) and del Hoyo et al. (2020) ZMUK). Plumage comparisons were made by directly com- “Tibet, central and northeastern China, the eastern half of paring museum specimens collected at the same time of the Mongolia and Transbaikalia”. The distribution in north- year, in comparable plumage and state of wear; plumage dif- ern Mongolia, northeastern China and Transbaikalia given ferences were only evaluated by eye, not by e.g. spectropho- by del Hoyo and Collar (2016) and del Hoyo et al. (2020) tometry. Measurements were taken by P.A. in the AMNH, is inhabited by C. brachydactyla according to the other ZMUC and UMMZ: wing length (with a ruler, wing fat- authors, whereas the eastern limit of C. brachydactyla is tened and stretched), tail length (with a ruler inserted under in west Mongolia according to del Hoyo and Collar (2016) the undertail-coverts) and bill length (with calipers, to base and del Hoyo et al. (2020). It is surprising that neither Cle- of skull). Wing and tail lengths were measured to the near- ments et al. (2019) nor Gill et al. (2020) include Mongolia est 0.5 mm, whereas bill length was measured to the nearest in the range of C. dukhunensis as the single breeding period 0.1 mm. sample of this taxon that was included in the phylogenetic To quantify diferences in song between C. dukhunensis analyses that form the basis for the recognition of C. dukhun- and C. brachydactyla, we analysed 8 high-quality record- ensis as a distinct species (Alström et al. 2013; Stervander ings of songs of C. dukhunensis and 24 recordings of et al. 2016) was collected in eastern Mongolia. Moreover, songs of C. brachydactyla, C. b. rubiginosa from Morocco neither Christidis (2018), Shirihai and Svensson (2018), Cle- (6); C. b. brachydactyla from Spain (3), France (2) and ments et al. (2019), Gill et al. (2020), del Hoyo and Collar Italy (3); and C. b. longipennis from Kazakhstan (5) and (2016) nor del Hoyo et al. (2020) appear to have noted that Xinjiang Province, China (5), i.e. eight C. dukhunensis Stervander et al. (2016) suggested that this complex does not and 6–10 of each of three additional subspecies, all difer- breed at all on the Tibetan Plateau. ent individuals (Supplementary Table S1; all new record- As the recently recognised species Calandrella dukhun- ings uploaded to Macaulay Library, Cornell University: ensis is extremely poorly known in most respects, with much https ://www.macau layli brary .org). Sonograms were cre- conficting information in the literature on its distribution, ated in Raven Pro 1.5 (Cornell Lab of Ornithology, USA), and with the recent elevation to species rank resting solely with the following settings: Window size = 256 samples; on molecular evidence, we here describe the morphology, window type = Hann; 3 dB flter bandwidth = 270 Hz; moult, breeding distribution, breeding habitat, vocalisations, overlap = 50%; size = 2.67 ms; DFT size = 256 samples; 1 3 Journal of Ornithology (2021) 162:165–177 167 spacing = 188 Hz. The contrast was kept fxed at 93, and xeno-canto (https://xeno-canto .org ) and own sound record- brightness was adjusted manually for each recording to ings (in total c. 15 C. dukhunensis and c. 100 C. brachy- give a good visual separation between the diferent song dactyla). As the song of C. dukhunensis is not divided elements and the background noise. The following vari- into strophes with pauses between (see Results) to produce ables were measured for each song strophe (Fig. 1): Dura- comparable measurements of the above variables (except tion, Minimum Frequency, Maximum Frequency, Band- pauses) we took measurements of C. dukhunensis songs width, Peak Frequency (the frequency associated with the from sequences of continuous song that corresponded to maximum energy) and the duration of Pauses between the mean value of the strophe length in C.
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