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Morphological and molecular investigations of Gagea (Liliaceae) in southeastern Kazakhstan with special reference to putative altitudinal hybrid zones

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Abstract

The most important center of speciation in the genus Gagea is thought to be in Central Asia. Here, we focus on species diversity in southeastern Kazakhstan (around Almaty, Ili-Alatau range of the Western Tian-Shan mountains). We studied an elevational transect, reaching from lowland steppes to the alpine zone (500–2750 m a. s. l.), and carried out detailed morphological and molecular investigations for populations of Gagea spp. Nine species were identified in different altitudinal zones; one of these (Gagea almaatensis) is described as new to science. We could detect two altitudinal contact zones between closely related species: G. filiformis and G. granulosa (sect. Minimae), and G. almaatensis and G. kuraminica (sect. Gagea). Morphological and molecular investigations (ITS data and cpDNA networks) indicate ongoing hybridization of co-occurring G. filiformis into G. granulosa and putative bidirectional hybridization events between G. almaatensis and G. kuraminica.

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Acknowledgments

For supporting field work and a research stay of S. Beisenova in Germany in the frame of its CabNet program, we wish to thank the German Academic Exchange Service (DAAD). Saltanat Beisenova wants to thank as well the authorities of N. I. Gumilyev Natl. Univ., Astana, for travel support and mentoring, in particular Prof. I. Bersimbayev. For providing further plant material, we would like to thank Jānis Rukšāns. We would like to thank the anonymous reviewers for helpful comments.

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Authors and Affiliations

  1. Institute of Biology, Martin-Luther-University of Halle-Wittenberg, Weinbergweg 10, 06120, Halle/Saale, Germany

    Angela Peterson

  2. Leibniz Institute of Plant Genetics and Crop Plant Research (IPK), 06466, Gatersleben, Germany

    Doerte Harpke

  3. Herbarium, Komarov Botanical Institute of the Russian Academy of Sciences, Prof. Popov Str. 2, Saint Petersburg, Russia, 197376

    Igor G. Levichev

  4. L.N. Gumilyov Eurasian National University, 5 Muneitpassov Str., Astana, Kazakhstan, 010008

    Saltanat Beisenova

  5. Institute of Botany and Landscape Ecology, Ernst Moritz Arndt University Greifswald, Soldmannstr. 15, 17487, Greifswald, Germany

    Martin Schnittler

  6. State Office for Environmental Protection of Saxony-Anhalt, Reideburger Str. 47, 06116, Halle/Saale, Germany

    Jens Peterson

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  1. Angela Peterson
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  2. Doerte Harpke
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Correspondence to Angela Peterson.

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Appendices

Appendix

Voucher information and GenBank accession numbers of the sequences used in this study. Taxon: place, date, collector, collection number: voucher number: EMBL/NCBI accession numbers: trnL–trnF IGS, psbA–trnH IGS, ITS1 + 5.8S rRNA + ITS2 (#sequences taken from EMBL/NCBI;—denotes lacking sequences). Information of collection dates, collectors, and collection numbers are only given for the material sampled in the Ili-Alatau range during the field excursion in 2013.

Gagea afghanica A.Terracc.: Iran, Province Khorasan: 52/04DNALevichevLE: HAL101785: –, –, AM087953. G. aipetriensis Levichev: Ukraine, Crimea: 15/04DNALevichevLE, HAL01801: AJ970178, AM049259, AM087955. G. almaatensis Levichev, A.Peterson & J.Peterson sp. nov.: (1) Kazakhstan, Ili-Alatau range, Almaty, 22 km S Almaty, Lake Bol. Almatinskoje, 2557 m a. s. l., 43°03′09″N, 76°59′23″E, 23 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP43a-2013: HAL119703, LE01010036: LN874781, LN874710, LN874873. (2) Kazakhstan, Ili-Alatau range, Almaty, 22 km S Almaty, Lake Bol. Almatinskoje, 2557 m a. s. l., 43°03′09″N, 76°59′23″E, 23 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP43b-2013: HAL119702: LN874782, LN874711, LN874874. (3) Kazakhstan, Ili-Alatau range, Almaty, 1.2 km NNW Shymbulak ski ressort, 1994 m a. s. l., 43°08′24″N, 77°04′25″E, 21 April 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP44a-2013: HAL119714: LN874783, LN874712, LN874875. (4) Kazakhstan, Ili-Alatau range, Almaty, 1.9 km S Shymbulak ski ressort, 2437 m a. s. l., 43°06′44″N, 77°04′26″E, 23 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP32-2013: HAL119695: LN874784, LN874713, LN874876. G. almaatensis × kuraminica: Kazakhstan: Ili-Alatau range: Almaty, 1.2 km NNW Shymbulak ski ressort, 1994 m a. s. l., 43°08′24″N, 77°04′25″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP24b-2013: HAL119688: LN874785, LN874714, LN874877. G. altaica Schischk. & Sumnev.: Kazakhstan, 51/04DNALevichevLE, HAL101786: –, –, AM162670. G. ancestralis Levichev: Russia, Republic Altai: 144/10DNALevichevLE: FR690249, FR690844, FR691052. G. angelae Levichev & Schnittler: (1) China, Xinjiang, Tian-Shan: LE00002135: FR690259, FR691028, FR690104. (2) China, Xinjiang, Tian-Shan: HAL108473: FR690266, FR691035, FR690111. (3) China, Xinjiang, Tian-Shan: HAL108477: FR690262, FR691031, –. G. artemczukii Krasnova: Ukraine, Crimea: 19/04DNALevichevLE, HAL101481: AM180470, AM409339, AM409331. G. bulbifera (Pall.) Salisb.: (1) China, Xinjiang, Urumqi: HAL107953: –, –, FR689753. (2) Russia, District Astrakhan: 2/04DNALevichevLE, HAL101840: –, –, AM162669. (3) Kazakhstan, Almaty, 14 km S Kapchagay, 521 m a. s. l., 43°44′24″N, 77°01′16″E, 19 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP11-2013: HAL118024: –, –, LN874788. (4) Kazakhstan, Almaty, 14 km S Kapchagay, 521 m a. s. l., 43°44′18″N, 77°00′43″E, 19 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP12-2013: HAL118026: –, –, LN874789. (5) Kazakhstan, Ili-Alatau range, Almaty, 2.1 km NE Medeu statium, 2003 m a. s. l., 43°10′12″N, 77°04′42″E, 20 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP18-2013: HAL118081: –, –, LN874790. (6) Kazakhstan, Ili-Alatau range, Almaty, 2.1 km NE Medeu statium, 2003 m a. s. l., 43°10′12″N, 77°04′42″E, 20 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler AP19a-2013: HAL119676, LE01010034: –, –, LN874791. (7) Kazakhstan, Ili-Alatau range, Almaty, 2.1 km NE Medeu statium, 2003 m a. s. l., 43°10′12″N, 77°04′42″E, 20 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP19b-2013: HAL119675: –, –, LN874792. (8) Kazakhstan, Ili-Alatau range, Almaty, 28 km W Almaty, small hills S Turar, 790 m a. s. l., 43°14′58″N, 76°32′13″E, 26 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP55-2013: HAL118028, LE01010035: –, –, LN874793. (9) Kazakhstan, Ili-Alatau range, Almaty, 29 km W Almaty, near settlement Uschkonyr, 864 m a. s. l., 43°13′15″N, 76°33′10″E, 26 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP57-2013: HAL118030: –, –, LN874794. G. capusii A.Terracc.: Uzbekistan: 24/04DNALevichevLE, HAL101839: AJ969123, AM085143, AM422455. G. circumplexa Vved.: Uzbekistan: 30/04DNALevichevLE, HAL101795: –, –, AM265529. G. confusa A.Terracc.: (1) Iran, Teheran: 13/04DNALevichevLE, HAL101803: AJ890369, AJ973173, AM087949. (2) #Iran: Zarrei&Zarrei35266(TUH)(Kew 23137): EU912267, EU939238, EU912040. (3) #Iran: TUH-EBOT.EXP.35712(TUH)(Kew 23169): EU912268, EU939239, EU912041. G. davlianidzeae Levichev: (1) China, Xinjiang, Tian-Shan: HAL107948: FR690236, FR690831, FR689755. (2) China, Xinjiang, Tian-Shan: HAL107943: FR690240, FR690835, FR689759. (3) China, Xinjiang, Tian-Shan: HAL107941: FR690241, FR690836, FR689760. (4) China, Xinjiang, Tian-Shan: HAL107946: FR690237, FR690832, –. (5) China, Xinjiang, Tian-Shan: HAL107939: FR690242, FR690837, –. G. divaricata Regel: (1) Kazakhstan, Almaty, 14 km S Kapchagay, 520 m a. s. l., 43°44′28″N, 76°01′20″E, 19 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP9a-2013: HAL118078, LE01010032: –, –, LN874795. (2) Kazakhstan: Almaty, 14 km S Kapchagay, 520 m a. s. l., 43°44′28″N, 76°01′20″E, 19 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP9b-2013: HAL118079: –, –, LN874796. (3) Kazakhstan, Kapchagay: 146/10DNALevichevLE: –, –, LN874797. G. dschungarica Regel: Kyrgyzstan: 14/04DNALevichevLE, HAL101802: –, –, AM087952. G. erubescens (Besser) Besser: Ukraine: 25/04DNALevichevLE, G-103LE, HAL103861: AM180469, AM238516, AM493953. G. fedtschenkoana Pascher: (1) Kazakhstan, Astana, mown lawns with steppe remnants near the administration building of the Gumilev University, Republican Ave, 360 m a. s. l., 51°10′52″N, 71°25′23″E: HAL117627: LN874767, LN874696, LN874849. G. cf. fedtschenkoana: (2) Kazakhstan, Tenis-Korgalshinskii nature reserve ca. 115 km WSW Astana, environs of the station 50°28′38″N, 250 m a. s. l., 69°32′44″E: HAL117626, LE01010043: LN874768, LN874697, LN874850. (3) Kazakhstan, Tenis-Korgalshinskii nature reserve ca. 115 km WSW Astana, steppe 2 km N of the station, 260 m a. s. l., 50°29′45″N, 69°32′48″E: HAL117628, LE01010044: LN874769, LN874698, LN874851. G. filiformis (Ledeb.) Kar. & Kir.: (1) Western Tian-Shan: 72/09DNALevichevLE: FR690250, FR690845, FR689768. (2) Kazakhstan: 12/04DNALevichevLE, HAL101804: AM084904, AM161459, AM180457. (3) Kazakhstan, Ili-Alatau foothills, Almaty, 5 km SE Almaty, Medeu valley, 1192 m a. s. l., 43°11′15″N, 77°00′07″E, 18 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP7b-2013: HAL118049: LN874716, LN874645, LN874809. (4) Kazakhstan: Ili-Alatau range: Almaty, 1.1 km NE Medeu statium, 1821 m a. s. l., 43°09′60″N, 77°04′28″, 20 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP14a-2013: HAL118050: LN874717, LN874646, LN874810. (5) Kazakhstan, Ili-Alatau range, Almaty, 1.1 km NE Medeu statium, 1821 m a. s. l., 43°09′60″N, 77°04′28″, 20 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP14b-2013: HAL118051: LN874718, LN874647, LN874811. (6) Kazakhstan, Ili-Alatau range, Almaty, 1.7 km NW Shymbulak ski ressort, 1941 m a. s. l., 43°08′30″N, 77°03′51″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP20a-2013: HAL118057, LE01010046: LN874719, LN874648, LN874812. (7) Kazakhstan, Ili-Alatau range, Almaty, 1.7 km NW Shymbulak ski ressort, 1941 m a. s. l., 43°08′30″N, 77°03′51″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler AP20b-2013: HAL118055: LN874720, LN874649, LN874813. (8) Kazakhstan, Ili-Alatau range, Almaty, 1.7 km NW Shymbulak ski ressort, 1941 m a. s. l., 43°08′30″N, 77°03′51″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP20c-2013: HAL118054: LN874721, LN874650, LN874814. (9) Kazakhstan, Ili-Alatau range, Almaty, 1.7 km NW Shymbulak ski ressort, 1941 m a. s. l., 43°08′30″N, 77°03′51″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP20d-2013: HAL118053: LN874722, LN874651, LN874815. (10) Kazakhstan, Ili-Alatau range, Almaty, 1.9 km S Shymbulak ski ressort, 2437 m a. s. l., 43°06′44″N, 77°04′06″, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler AP30a-2013: HAL119715: LN874723, LN874652, LN874816. (11) Kazakhstan, Ili-Alatau range, Almaty, 1.9 km S Shymbulak ski ressort, 2437 m a. s. l., 43°06′44″N, 77°04′06″, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP30b-2013: HAL118062: LN874724, LN874653, LN874817. (12) Kazakhstan, Ili-Alatau range, Almaty, 1.5 km SSW Shymbulak ski ressort, 2380 m a. s. l., 43°06′59″N, 77°04′26″E, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP33a-2013: HAL118065: LN874725, LN874654, LN874818. (13) Kazakhstan, Ili-Alatau range: Almaty, 1.5 km SSW Shymbulak ski ressort, 2380 m a. s. l., 43°06′59″N, 77°04′26″E, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP33b-2013: HAL118064: LN874726, LN874655, LN874819. (14) Kazakhstan, Ili-Alatau range, Almaty, 1.5 km SSW Shymbulak ski ressort, 2380 m a. s. l., 43°06′59″N, 77°04′26″E, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP34-2013: HAL118063: LN874727, LN874656, LN874820. (15) Kazakhstan, Ili-Alatau range, Almaty, 1.5 km NW Medeu stadium, 1498 m a. s. l., 43°10′09″N, 77°02′25E, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP38a-2013: HAL118067: LN874728, LN874657, LN874821. (16) Kazakhstan, Ili-Alatau range, Almaty, 1.5 km NW Medeu stadium, 1498 m a. s. l., 43°10′09″N, 77°02′25″E, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP38b-2013: HAL118068: LN874729, LN874658, LN874822. (17) Kazakhstan, Ili-Alatau range, Almaty, 22 km S Almaty, Lake Bol. Almatinskoje, 2733 m a. s. l., 43°03′22″N, 76°58′05″E, 23 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP42-2013: HAL118069: LN874730, LN874659, LN874823. (18) Kazakhstan, Ili-Alatau foothills, Almaty, 4 km SE Almaty, around TV tower, 1099 m a. s. l., 43°13′36″N, 76°58′26″E, 25 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP51-2013: HAL118070: LN874731, LN874660, LN874824. (19) Kazakhstan, Ili-Alatau foothills, Almaty, 4 km SE Almaty, around TV tower, 1089 m a. s. l., 43°13′50″N, 76°58′29″E, 25 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP52-2013: HAL118073: LN874732, LN874661, LN874825. (20) Kazakhstan: Ili-Alatau foothills: Almaty, 3 km NW Medeu stadium, 1301 m a. s. l., 43°10′48″N, 77°00′43″E, 27 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP59a-2013: HAL118074: LN874733, LN874662, LN874826. (21) Kazakhstan: Ili-Alatau foothills, Almaty, 3 km NW Medeu stadium, 1301 m a. s. l., 43°10′48″N, 77°00′43″E, 27 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP59b-2013: HAL118075: LN874734, LN874663, LN874827. (22) Kazakhstan: Ili-Alatau foothills: Almaty, 3 km NW Medeu stadium, 1301 m a. s. l., 43°10′48″N, 77°00′43″E, 27 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP59c-2013: HAL118076: LN874735, LN874664, LN874828. (23) Kazakhstan: Ili-Alatau range: Almaty, 1 km SSE Medeu stadium, 1875 m a. s. l., 43°09′06″N, 77°04′01″, 29 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP60-2013: HAL118077: LN874736 LN874665, LN874829. (24) Kazakhstan, Ili-Alatau range, Almaty, 1.2 km NNW Shymbulak ski ressort, 1994 m a. s. l., 43°08′24″N, 77°04′25″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP25-2013: HAL118059: LN874737, LN874666, LN874830. (25) Kazakhstan, Mountains over Almaty, at Observatory: DNAIPK12KZ-123: –, –, LN874831. (26) Kazakhstan: Sharyn Canyon: HAL115984: LN874738, LN874667, LN874832. (27) Kazakhstan, Mramornij (Marble) pass, W end of Altai: DNA IPK12KZ-102: –, –, LN874833. (28) Kazakhstan, near Kegen Pass, HAL116095: LN874739, LN874668, LN874834. (29) Kazakhstan, Ili-Alatau range, Almaty, 1.7 km NW Shymbulak ski ressort, 1941 m a. s. l., 43°8′30″N, 77°03′51″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP23-2013: HAL118058: LN874740, LN874669, –. (30) Kazakhstan, Ili-Alatau foothills: Almaty, 5 km SE Almaty, Medeu valley, 1192 m a. s. l. 43°11′15″N, 77°00′07″, 18 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP7c-2013: HAL118039: LN874753, LN874682, LN874847. G. graeca (L.) Irmisch.: Greece, Crete: HAL105185: –, –, AM939649. G. graminifolia Vved.: Kyrgyzstan, 114/09DNALevichevLE: –, –, FR689769. G. granulosa Turcz.: (1) –: 11a/04DNALevichevLE, HAL101805: –, –, AM265533. (2) Russia, Sibiria: 141/09DNALevichevLE: FR690253, FR690848, –. (3) Ukraine, Poltava: 100/09DNALevichevLE: FR690252, FR690847, –. (4) Kazakhstan, Tian-Shan: 11b/04DNALevichevLE, HAL101836: AM180463, AM238517, AM287278. (5) Kazakhstan, Ili-Alatau foothills, Almaty, 862 m a. s. l., 43°13′34″N, 76°54′49″E, 18 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP4-2013: HAL118037: LN874741, LN874670, LN874835. (6) Kazakhstan, Ili-Alatau foothills, Almaty, 839 m a. s. l., 43°14′57″N, 76°56′42″E, 17 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP1a-2013: HAL118035, LE01010049: LN874742, LN874671, LN874836. (7) Kazakhstan, Ili-Alatau foothills, Almaty, 839 m a. s. l., 43°14′57″N, 76°56′42″E, 17 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP1b-2013: HAL118033: LN874743, LN874672, LN874837. (8) Kazakhstan, Ili-Alatau foothills, Almaty, 5 km SE Almaty, Medeu valley, 1192 m a. s. l., 43°11′15″N, 77°00′07″E, 18 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP7a-2013: HAL118038: LN874744, LN874673, LN874838. (9) Kazakhstan, Ili-Alatau range, Almaty, 300 m NW Medeu statium, 1721 m a. s. l., 43°09′37″N, 77°03′33″E, 20 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP13-2013: HAL118040: LN874745, LN874674, LN874839. (10) Kazakhstan, Ili-Alatau range, 1.7 km NW Shymbulak ski ressort, 1941 m a. s. l., 43°08′30″N, 77°03′51″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP21-2013: HAL118042: LN874746, LN874675, LN874840. (11) Kazakhstan, Ili-Alatau range, Almaty, 1.2 km NNW Shymbulak ski ressort, 1994 m a. s. l., 43°08′24″N, 77°04′25″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP28-2013: HAL118043: LN874747, LN874676, LN874841. (12) Kazakhstan, Ili-Alatau range, Almaty, 1.7 km NW Shymbulak ski ressort, 1941 m a. s. l., 43°08′30″N, 77°03′51″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP45a-2013: HAL118046: LN874748, LN874677, LN874842. (13) Kazakhstan, Ili-Alatau range, Almaty, 1.7 km NW Shymbulak ski ressort, 1941 m a. s. l., 43°08′30″N, 77°03′51″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP45b-2013: HAL118045: LN874749, LN874678, LN874843. (14) Kazakhstan, Ili-Alatau range, Almaty, 1.7 km NW Shymbulak ski ressort, 1941 m a. s. l., 43°08′30″N, 77°03′51″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP45c-2013: HAL118044: LN874750, LN874679, LN874844. (15) Kazakhstan, Ili-Alatau foothills, Almaty, 3 km NW Medeu stadium, 1301 m a. s. l., 43°10′48″N, 77°00′43″E, 27 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP58a-2013: HAL118048: LN874751, LN874680, LN874845. (16) Kazakhstan, Ili-Alatau foothills, Almaty, 3 km NW Medeu stadium, 1301 m a. s. l., 43°10′48″N, 77°00′43″E, 27 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP58b-2013: HAL118047: LN874752, LN874681, LN874846. (17) Kazakhstan, near Panteleimonovka vil.,: DNA12KZ-117: –, –, LN874848. G. helenae Grossh: Russia, Dagestan: 22/04DNALevichevLE, HAL101798: AJ969120, AM161461, AM265531. G. huochengensis Levichev: (1) China, Xinjiang, Tian-Shan: HAL108466: FR690269, FR691038, FR690114. (2) Kazakhstan: 132/09DNALevichevLE, HAL107949: FR690274, FR691042, FR690118. (3) China, Xinjiang, Tian-Shan: HAL108468: FR690272, FR691041, –. (4) China, Xinjiang, Tian-Shan: HAL108467: FR690271, FR691040, –. G. jensii Levichev & Schnittler: China, Xinjiang: HAL108482: FR690256, FR691025, FR690101. G. kuraminica Levichev (1) Kazakhstan, Ili-Alatau foothills, Almaty, 5 km SE Almaty, Medeu valley, 1192 m a. s. l., 43°11′15″N, 77°00′07″E, 18 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP6-2013: HAL119683, LE01010038: LN874770, LN874699, LN874852. (2) Kazakhstan, Ili-Alatau range, Almaty, 1.5 km NE Medeu statium, 1920 m a. s. l., 43°10′02″N, 77°04′36″E, 20 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP16-2013: HAL119685, LE01010050: LN874771, LN874700, LN874853. (3) Kazakhstan, Ili-Alatau range, Almaty, 1.5 km NW Medeu stadium, 1498 m a. s. l., 43°10′09″N, 77°02′25″E, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP49-2013: HAL119704, LE01010042: LN874772, LN874701, LN874854. (4) Kazakhstan, Ili-Alatau range, Almaty, 1.2 km NNW Shymbulak ski ressort, 1994 m a. s. l., 43°08′24″N, 77°04′25″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP24a-2013: HAL119687: LN874773, LN874702, LN874855. (5) Kazakhstan, Ili-Alatau range, Almaty, 1.2 km NNW Shymbulak ski ressort, 1994 m a. s. l., 43°08′24″N, 77°04′25″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP24c-2013: HAL119689: LN874774, LN874703, LN874856. (6) Kazakhstan, Ili-Alatau range, Almaty, 1.5 km SSW Shymbulak ski ressort, 2380 m a. s. l., 43°06′59″N, 77°04′26″E, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP36c-2013: HAL119697: LN874775, LN874704, LN874857. (7) Kazakhstan, Ili-Alatau range, Almaty, 1.2 km NNW Shymbulak ski ressort, 1994 m a. s. l., 43°08′24″N, 77°04′25″E, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP44b-2013: HAL119713: LN874776, LN874705, LN874858. (8) Kazakhstan, Ili-Alatau range, Almaty, 1.5 km SSW Shymbulak ski ressort, 2380 m a. s. l., 43°06′59″N, 77°04′26″E, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP37-2013: HAL119700, LE01010038: LN874777, LN874706, LN874859. (9) Kazakhstan, Ili-Alatau range, Almaty, 1.9 km S Shymbulak ski ressort, 2437 m a. s. l., 43°06′44″N, 77°04′26″E, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP29a-2013: HAL119693, LE01010040: LN874778, LN874707, LN874860. (10) Kazakhstan, Ili-Alatau range, Almaty, 1.9 km S Shymbulak ski ressort, 2437 m a. s. l., 43°06′44″N, 77°04′26″, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP29b-2013: HAL119692: LN874779 LN874708, LN874861. (11) Uzbekistan, Tian-Shan, Kurama range, 2500 m a. s. l.,: 151/14DNALevichevLE (holotype): LN874780, LN874709, LN874862. G. kuraminica × almaatensis Kazakhstan, Ili-Alatau range, Almaty, 22 km S Almaty, Lake Bol. Almatinskoje, 2733 m a. s. l., 43°03′22″N, 76°58′05″E, 23 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP40-2013: HAL119701: LN874786, LN874715, LN874878. G. liotardii (Sternberg) Schult. et Schult.f. (= G. fragifera (Vill.) Ehr.Bayer & G.López): (1) China, Xinjiang, Tian-Shan: HAL107898: –, –, FN868162. (2) Ukraine, Crimea: 29a/04DNALevichevLE, HAL101797: –, –, AM265532. (3) Italy: CLU12693: –, –, AM422468. (4) Kazakhstan: 29b/04DNALevichev LE, HAL101796: –, –, AM180455. (5) Kazakhstan, Tenis-Korgalshinsky Nature Reserve, Astana: HAL117634: –, –, LN874799. (6) Kazakhstan, Ili-Alatau range, Almaty, 1.7 km NW Shymbulak ski ressort, 1941 m a. s. l., 43°08′30″N, 77°03′51″, 21 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP22-2013: HAL118013, LE01010041: –, –, LN874800. (7) Kazakhstan, Ili-Alatau range, Almaty, 1.9 km S Shymbulak ski ressort, 2437 m a. s. l., 43°06′44″N, 77°04′26″, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP31-2013: HAL118015: –, –, LN874801. (8) Kazakhstan, Ili-Alatau range, 1.5 km NW Medeu stadium, road to Medeu, 1498 m a. s. l., 43°10′09″N, 77°02′25″E, 22 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP39-2013: HAL118018: –, –, LN874802. (9) Kazakhstan, Ili-Alatau range, Lake Bol. Almatinskoje, 2733 m a. s. l., 43°03′22″N, 76°58′05″E, 23 Apr 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP41-2013: HAL118032: –, –, LN874803. (10) Kazakhstan, Mountains over Almaty: DNA IPKKZ124: –, –, LN874804. (11) Morocco: Marrakech: HAL116006: –, –, LN874805. G. lutea Ker Gawl.: (1) Germany, Saxony-Anhalt: –: AJ419165, AJ416368, AJ427544. (2) Russia, Caucasus: 16/04DNALevichevLE, HAL101800: AM110255, AM161456, AM265530. (3) Italy, Calabria CLU12696: AM265606, AM409354, AM287282. (4) #Iran: Zarrei35285(TUH)(Kew 23148): EU912281, EU939250, EU912052. G. marchica Henker, Kiesew., U.Raabe & Rätzel: Germany, Alt-Zeschdorf: HAL118006: LN874754, LN874683, LN874808. G. megapolitana Henker: Germany: HAL101644: AM084902, AM161455, AM931553. G. minima (L.) Ker Gawl.: (1) Italy, Calabria, Pollino Massif: CLU12698: AM283109, AM282996, AM287273. (2) Ukraine, Carpathians: 9/04DNALevichevLE, HAL101808: AM238539, AM238524, AM087948. (3) Russia, Republic Tatarstan, Nizhnyaya Kama: 10/04DNALevichevLE, HAL101806: AM180471, AM238519, AM180459. (4) Germany, Saxony-Anhalt: –: AJ419164, AJ416374, AJ427546. G. nakaiana Kitag.: Russia, Khabarovsk Territory: 17/04DNALevichev, 55LE, HAL101799: AM110256, AM161457, AM180454. G. neopopovii Golosk.: Eastern Kazakhstan, Za-Iliskii Altai: 120/09DNALevichevLE: –, –, FR689770. G. nigra L.Z.Shue, emend. Schnittler: (1) China, Xinjiang, Urumqi: HAL108464: FR690123, FR691010, FR690081. (2) China, Xinjiang, Urumqi: HAL108452: FR690135, FR691022, FR690095. (3) China, Xinjiang, Urumqi (N9714, Sichuan Univ., Chengdu): 134/10DNALevichevLE: FR690137, FR691024, –. (4) China, Xinjiang, Tian-Shan, Bogda-Shan, Tianchi Lake: HAL108459: FR690125, FR691012, FR690083. (5) China, Xinjiang, Tian-Shan, Bogda-Shan, Tianchi Lake: HAL108460: FR690128, FR691015, –. (6) China, Xinjiang, Tian-Shan, Bogda-Shan, Tianchi Lake: HAL108456: FR690129, FR691016, –. (7) China, Xinjiang, Tian-Shan, Bogda-Shan, Tianchi Lake: HAL108183: FR690131, FR691018, –. (8) China, Xinjiang, Tian-Shan, Bogda-Shan, Tianchi Lake: HAL108457: FR690132, FR691019, –. (9) China, Xinjiang, Tian-Shan, Bogda-Shan: HAL108184: FR690134, FR691021, –. G. ova Stapf: China, Xinjiang, Urumqi: HAL107951: –, –, FR689762. G. paczoskii (Zapal.) Grossh.: Italy: HAL104323: AM903072, AM903061, AM903051. G. pauciflora Turcz. ex Ledeb.: Mongolia, Ulan Bator: HAL070423: –, –, AM409330. G. podolica Schult. & Schult.f.: Ukraine: 21/04DNALevichevLE, HAL101832: AM084903, AM238525, AM409334. G. pomeranica Ruthe: (1) Germany, Saxony-Anhalt HAL095842: AJ419167, AJ416375, AJ427543. (2) Germany, Mecklenburg-Western Pomeranica: HAL095846: LN874787, AJ429194, –. G. pratensis (Pers.) Dumort.: (1) Germany, Saxony-Anhalt: HAL095847: AJ419162, AJ416372, AJ427542. (2) Italy: CLU12703, AM265607, AM903059, AM903047. G. pusilla (F.W.Schmidt) Sweet: cultivated: 18/04DNALevichevLE, HAL101831: AM180464, AM161458, AM422453. G. reticulata (Pall.) Schult. et Schult.f.: Israel, Central Negev: Z34650: –, –, AM087954. G. rubicunda Meinsh. emend. Levichev: Estonia: 26/04DNALevichevLE, HAL103855: AM238541, AM409338, AM493954. G. rufidula Levichev: (1) China, Xinjiang, Tian-Shan: HAL107929: –, –, FR689764. (2) Kyrgyzstan: 149/09DNALevichevLE: –, –, FR689767. G. serotina (L.) Ker Gawl. (=Lloydia serotina (L.) Reichenb.): (1) Kazakhstan: 45a/04DNALevichevLE, HAL101789: –, –, AM087956. (2) Russia, Karachaevo-Cherkessian Rep.: HAL117624: –, –, LN874806. (3) USA, Alaska, Alaska range: HAL117490: –, –, LN874807. G. shmakoviana Levichev: Russia, Altai: 23/04DNALevichevLE, HAL101830: AM265520, AM287265, AM422454. G. spathacea (Hayne) Salisb.: Germany, Saxony-Anhalt: HAL095844: –, –, AJ427541. G. stipitata Merckl. ex Bunge: Iran, Province Khorasan: 49/04DNALevichevLE, HAL101828: –, –, AM409336. G. tenera Pascher: (1) Kyrgyzstan: HAL101843: –, –, AM422460. (2) Iran: Zarrei & Golzarian 35256(TUH)(Kew 23152): –, –, EU912074. (3) Kazakhstan, Ili-Alatau foothills, Almaty, 831 m a. s. l., 43°15′0″N, 76°56′41″E, 18 April 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP3-2013: HAL118080, LE01010033: –, –, LN874798. G. terraccianoana Pascher: (1) Russia: 55/04DNALevichevLE, HAL103853: AM287280, AM287268, AM493955. (2) Mongolia: Bogd-Ul Mountains: HAL070426: AJ890367, AJ973169, –. G. tisoniana Peruzzi, Bartolucci, Frignani & Minutillo: Italy: CLU14920: AM409358, AM409349, AM422466. G. transversalis Steven: Ukraine, Crimea: 56/04DNALevichevLE, 365LE, HAL101783: AJ890370, AJ973167, AM162671. G. turkestanica Pascher: (1) Kyrgyzstan: 20/04DNALevichevLE, HAL103850: AM238534, AM287267, –. G. cf. turkestanica (2) Kazakhstan, Ili-Alatau range, Almaty, 28 km W Almaty, small hills S Turar, 790 m a. s. l., 43°14′58″N, 76°32′13″E, 26 April 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP54a-2013: HAL119707, LE01010039: LN874755, LN874684, LN874863. (3) Kazakhstan: Ili-Alatau range: Almaty, 28 km W Almaty, small hills S Turar, 43°14′58″N, 76°32′13″E, 790 m a. s. l., 26 April 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP54b-2013: HAL119710: LN874756, LN874685, LN874864. (4) Kazakhstan, Ili-Alatau foothills, Almaty, Panfilov park, 831 m a. s. l., 43°15′00″N, 76°56′41″E, 24 April 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP46-2013: HAL118082: LN874757, LN874686, LN874865. (5) Kazakhstan, Almaty: 130/09DNALevichevLE: LN874758, LN874687, LN874866. (6) Kazakhstan, Jezkazgan: 131/09DNALevichevLE: LN874759, LN874688, –. (7) Kazakhstan, Almaty, 950 m a. s. l.: 119/09DNALevichevLE: LN874760, LN874689, –. (8) Kazakhstan, Ili-Alatau foothills: Almaty, 831 m a. s. l., 43°15′00″N, 76°56′41″E, 18 April 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP2a-2013: HAL119678: LN874761, LN874690, LN874867. (9) Kazakhstan, Ili-Alatau foothills, Almaty, 831 m a. s. l., 43°15′00″N, 76°56′41″E, 18 April 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP2a-2013: HAL119679: LN874762, LN874691, LN874868. (10) Kazakhstan, Ili-Alatau foothills, Almaty, 862 m a. s. l., 43°13′34″N, 76°54′49″E, 18 April 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP2a-2013: HAL119680: LN874763, LN874692, LN874869. (11) Kazakhstan, Ili-Alatau foothills, Almaty, 4 km SE Almaty, park around the TV tower, 1119 m a. s. l., 43°13′49″N, 76°58′38″E, 25 April 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP53-2013: HAL119708: LN874764, LN874693, LN874870. (12) Kazakhstan, Ili-Alatau foothills, Almaty, 4 km SE Almaty, around TV tower, 1099 m a. s. l., 43°13′36″N, 76°58′26″E, 25 April 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP50-2013: HAL119705: LN874765, LN874694, LN874871. (13) Kazakhstan, Ili-Alatau range, Almaty, 29 km W Almaty, near settlement Uschkonyr, 864 m a. s. l., 43°13′15″N, 76°33′10″E, 26 April 2013, A. and J.Peterson, S.Beisenova, M.Schnittler, AP56-2013: HAL119711: LN874766, LN874695, LN874872. G. uliginosa Siehe & Pascher: #Iran: Zarrei et al. TUH-EBOT.EXP.35304,(TUH)(Kew 23153): –, –, EU912089. G. vegeta Vved.: Uzbekistan: 32/04DNALevichev, 339aLE, HAL103691: –, –, AM287275. G. villosa (Bieb.) Sweet: Germany, Saxony-Anhalt: –: –, AJ427545. G. xiphoidea Levichev: (1) Russia, Republic Altai: 71/09DNALevichevLE: FR690138, FR691044, FR690097. (2) Russia, Republic Altai, district Kosch-Agaczensis: 102/09DNALevichevLE: FR690139, FR691045, FR690098. G. cf. xiphoidea (3) China, Xinjiang, Tian-Shan: HAL108479: FR690140, FR691046, FR690099. Tulipa cretica Boiss. & Heldr.: Greece, Crete: HAL099934: –, –, AM180461.

Information on Electronic Supplementary Material

Online Resource 1 ITS alignment used for BI and resulting tree.

Online Resource 2 cpDNA alignment used for TCS of sect. Minimae.

Online Resource 3 cpDNA alignment used for TCS of sect. Gagea.

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Peterson, A., Harpke, D., Levichev, I.G. et al. Morphological and molecular investigations of Gagea (Liliaceae) in southeastern Kazakhstan with special reference to putative altitudinal hybrid zones. Plant Syst Evol 302, 985–1007 (2016). https://doi.org/10.1007/s00606-016-1313-7

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