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Phylogeography of the House Bunting complex: discordance between species limits and genetic markers

Journal of Ornithology volume 159, pages 47–61 (2018)Cite this article

Abstract

Emberiza striolata has traditionally been viewed as a single polytypic species, but recent taxonomic reviews of its morphology, vocalizations and genetic data have led to the recommendation that two species be recognized within this complex, the House Bunting Emberiza sahari in the west, and the Striolated Bunting Emberiza striolata (including the subspecies Emberiza striolata saturiator and Emberiza striolata ‘jebelmarrae’) in the east of the range. However, reportedly intermediate birds have been described from northern Darfur, Sudan, and the Ennedi Massif, northeast Chad, bringing the two-species hypothesis into question. Here, we attempt a more comprehensive geographic sampling of all taxa of E. sahari and E. striolata, and we report the results of an examination of purported intermediate specimens, as well as the re-examination of the long type series of the controversial taxon Emberiza striolata ‘jebelmarrae’ described from western Sudan. Our results confirm a marked east–west divergence in the House Bunting complex based on two mitochondrial DNA (mtDNA) markers, but surprisingly reveal that samples of E. striolata ‘jebelmarrae’ cluster with the western clade, whereas their morphology is clearly allied with that of other populations in the eastern clade. However, our analysis suggests that all of the specimens believed to be intermediate in morphology can be categorically placed with other examples of the western clade (E. sahari). At present, it is impossible to definitively explain the phylogenetic position of E. striolata ‘jebelmarrae’, but we hypothesize that the paraphyly in mtDNA is the result of introgression during the initial phase of secondary contact, and does not impinge on the decision to recognize two species. Complex spatio-temporal population patterns caused by late Pleistocene climate changes might have led to multiple secondary contact events between the eastern and western clades.

Zusammenfassung

Phylogeographie der Hausammern: ein Widerspruch zwischen Artgrenzen und der Verteilung der genetischen Vielfalt

Die Hausammer, Emberiza striolata, wurde traditionellerweise als eine polytypische Art betrachtet. Neuere taxonomische Revisionen basierend auf morphologischen, akustischen und genetischen Daten empfehlen dagegen eine Aufspaltung in zwei Arten, die Hausammer E. sahari im Westen und die Wüstenammer E. striolata (inklusive der Unterarten saturiator and ‘jebelmarrae’) im Osten des Verbreitungsgebietes. Aus der nördlichen Darfur-Region Sudans und dem Ennedi Massif im Nordosten Tschads sind aber scheinbar intermediäre Individuen beschrieben worden und lassen daher Zweifel an der Zwei-Arten-Hypothese aufkommen. In dieser Studie bearbeiten wir eine umfassendere geographische Stichprobe als bisherigen genetische Arbeiten und untersuchen zudem die mutmasslich intermediären Exemplare sowie die Typenserie des kontrovers diskutieren Taxons ‘jebelmarrae’, das aus dem Westen Sudans beschrieben wurde. Eine klare Ost-West Trennung innerhalb des gesamten Hausammer-Komplexes wird in unseren Analysen von zwei mitochondrialen Markern eindeutig unterstützt. Überraschenderweise finden sich die untersuchten Individuen von ‘jebelmarrae’ innerhalb der westlichen Gruppe (sahari), obwohl dieses Taxon morphologisch eindeutig der östlichen Gruppe (striolata) zugewiesen werden kann. Alle Exemplare mit vermeintlich intermediär ausgeprägten Merkmalen lassen sich aufgrund unserer Studie morphologisch eindeutig der westlichen Gruppe zuordnen. Die phylogenetische Position von ‘jebelmarrae’ bleibt weiterhin unklar. Die Paraphylie in den mitochondrialen Markern interpretieren wir als die Folge von Introgression zwischen der östlichen und der westlichen Gruppe während einer anfänglichen Phase von sekundärem Kontakt; sie widerspricht daher nicht der Anerkennung von zwei Arten. Klimatische Veränderungen während der letzten Phase des Pleistozäns führten vermutlich zu einer zeitlich und räumlich komplizierten Populationsgeschichte mit wiederholtem sekundärem Kontakt zwischen der westlichen und der östlichen Gruppe.

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Fig. 1

[Distribution areas are modified from BirdLife International and the Handbook of the Birds of the World (2016). Bird figures were taken with permission from del Hoyo et al. (2017)]

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Acknowledgments

We are grateful to the following museum curators for permission to examine and sample specimen material: Sylke Frahnert and Pascal Eckhoff (Museum für Naturkunde, Berlin), Till Töpfer (ZFMK, Bonn), Fritz Geller-Grimm (MWNH, Wiesbaden), and Mark Adams, Hein van Grouw and Robert Prŷs-Jones (NHMUK, Tring). Original research by G. M. K. and H. S. on the taxonomy of E. striolata involved visits to several additional museums, whose curatorial staff are acknowledged in Kirwan and Shirihai (2007). The Israeli samples were from feathers taken by H. S., who is grateful to Yosef Kiat of the Israeli Bird Ringing Center (Israel Ornithological Center, Society for the Protection of Nature in Israel) for enabling studies of Striated Buntings in the hand during the annual ringing programme at the Dead Sea. Another sample was taken from the tissue collection of the Steinhardt Museum of Natural History and National Research Center, Department of Zoology, Tel Aviv University, with the help of Daniel Berkowic and Dr Roi Dor. M. S. thanks Elisa Badia (Lynx), Mark Balman (BirdLife), Tom Burri and Chris Sherry for their help with Fig. 1. Heiko Schmaljohann is financed by the German Research Foundation (SCHM 2647/1-2).

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  1. Naturhistorischen Museum der Burgergemeinde Bern, Bernastrasse 15, 3005, Bern, Switzerland

    Manuel Schweizer

  2. c/o Ausserdorfstrasse 6, 8052, Zurich, Switzerland

    Hadoram Shirihai

  3. Institute of Avian Research “Vogelwarte Helgoland”, An der Vogelwarte 21, 26386, Wilhelmshaven, Germany

    Heiko Schmaljohann

  4. Field Museum of Natural History, 1400 South Lakeshore Drive, Chicago, IL, 60605, USA

    Guy M. Kirwan

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  1. Manuel Schweizer
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Correspondence to Manuel Schweizer.

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Schweizer, M., Shirihai, H., Schmaljohann, H. et al. Phylogeography of the House Bunting complex: discordance between species limits and genetic markers. J Ornithol 159, 47–61 (2018). https://doi.org/10.1007/s10336-017-1501-4

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