Gagea bulbifera (Pall.) Salisb. Ann. Bot.

Tekşen, Mehtap & Erkul, Seher Karaman, 2015, The synopsis of the genus Gagea (Liliaceae) in Turkey, Phytotaxa 230 (1), pp. 101-129 : 103-105

Gagea bulbifera (Pall.) Salisb. Ann. Bot. 2: 557 (1806)

≡ Ornithogalum bulbiferum Pall., Reise Russ. Reich. 2: 736 (1773) ≡ Stellaster bulbifer (Pall.) Kuntze, Revis. General. Pl. 2: 715 (1891) ≡ Hornungia bulbifera (Pall.) Rouy in Rouy & Fouc., Fl. France 12: 381 (1910) = Gagea bulbifera (Pall.) Salisb. var. major Besser ex Schult. & Schult.f. in Roem. & Schult., Syst. Veg. 7: 552 (1829) = Gagea polyphylla Steven, Bull. Soc. Imp. Naturalistes Moscou 30(2): 83 (1857) = Gagea bulbifera (Pall.) Salisb. var. nuda Regel, Trudy Imp. S. -Peterburgsk. Bot. Sada 6: 511 (1879) = Gagea bulbifera var. chersonica Lindemann, Fl. Cherson. 2: 228 (1882) = Gagea bulbifera (Pall.) Salisb. var. armena Miscz. Fl. Cauc. Crit. 2(4): 175 (1912) = Gagea bulbifera f. ebulbillifera Grossh., Trudy Obshch. Isp. Prir. Imp. Khar’kovsk. Univ. 49: 9 (1918).

Type (lectotype designated by Stroh 1937: 501): — KAZAKYSTAN. (Astrakhan), in aridis limosis ad Wolgam et Jaicum, Pallas s.n., Herb. Linn. 428.17 ( E!).

Specimens seen: — TURKEY. Ağrı: Patnos, N of Suphan mountain, Erkeçli village , 2300–2500 m, 26 June 2002, L. Behçet 6956 ( VANF!) ; Patnos , Top mountain, meadows, rocky slopes, 1850 m, 27 April 2007, H.684 ( VANF!) ; Kağızman , steppe, 15 April 1957, K. Karamanoğlu 264 ( ANK!) ; Bitlis: Tatvan, Ortatepe, above Tatvan , alpine steppe, c. 2600 m, 19 May 1972, H. Peşmen 2735 ( HUB!) ; Kurudere village , rocky slopes, 1200 m, 23 April 2002, A. A. 2075 ( VANF!) ; S slopes Alacabük mountain, above Kuskunkıran pass, steppe, 2500–3000 m, 27 April 2001, F. Özgökçe 9258 ( VANF!) ; Adilcevaz, N of Bahçedeme village , 2100 m, 10 May 1988, L. Behçet 670 ( VANF!) ; Erzincan: Çağlayan, Çukur arable field, Munzur mountains , 1350–1600 m, 23 April 1980, Ş. Yıldırımlı 2587 ( ANK!) ; Erzurum: Horasan-Kars, 2 km from Horasan , steppe, 1539 m, 26 April 2011, M. Tekşen 2782 & S. Karaman ( AKSU!) ; Tahir mountain, near Aktaş , 21 May 1976, T. Baytop 34909 ( E!) ; Gümüşhane: Gümüşhane-Bayburt road, 5 May 1960, Stainton 8350 ( E!) ; Çiflik Village on University Farm , 14 May 1967, J. G. Ross s.n. ( E!) ; Erzurum SW, Atatürk University , meadows, 1900 m, 8 May 1969, S. Özyurt, N. Balta 23589 ( ISTF!) ; on Atatürk University Farm , meadows, 1850 m, 16 April 1970, S. Özyurt, N. Balta 24649 ( ISTF!) ; Atatürk University , meadows, 1890 m, 14 April 1970, S. Özyurt 24648 ( ISTF!) ; Palandöken mountains, E slopes of Sultanseki hills, chaierlift environment, stony meadows, 2200 m, 25 May 1970, S. Özyurt, N. Balta 24941 ( ISTF!) ; Horasan-Kars, 1 km beyond Horasan , steppe, 1535 m, 26 April 2011, M. Tekşen 2780 & S. Karaman ; Iğdır: 6 km from Doğubeyazıt to Iğdır, rocky slopes, 1540 m, 23 April 2002, A. A. Dönmez 10540 ( HUB!) ; Kars: Sarıkamış, Karakurt, Agop state, 1450–1600 m, 11 May 1980, O. Güneş 1534 ( HUB!) ; Kale , rock cracks, 1750 m, 2 June 1993, A. A. Dönmez 3236 & Metzger ( HUB!) ; Muş: Malazgirt , SE of Baklaya village, fields, 1657 m, 16 April 2006, LFM 58 ( VANF!) ; Malazgirt , Aktuzlu village, steppe, 1550 m, 7 April 2001, S.1193 ( VANF!) ; Malazgirt , steppe, 1531 m, 29 April 2006, LFM 129 ( VANF!) ; Van: Hoşap-Van, 6 km beyond Hoşap , steppe, 1957 m, 27 April 2011, M. Tekşen 2786 & S. Karaman ( GAZI!) ; Hoşap-Van, 6.5 km beyond Hoşap , steppe, 1947 m, 27 April 2011, M. Tekşen 2789 & S. Karaman ( AKSU!) ; Hoşap-Van, 12 km beyond Hoşap , steppe, 1968 m, 27 April 2011, M. Tekşen 2793 & S. Karaman ( AKSU!) ; Hoşap-Van, 12 km beyond Hoşap , rocky places, 1968 m, 27 April 2011, M. Tekşen 2794 & S. Karaman ( AKSU!) ; Hoşap-Van, 13 km beyond Hoşap, Zernek dam environment, steppe, wet places, 1961 m, 27 April 2011, M. Tekşen 2799 & S. Karaman ( AKSU!) ; Hoşap-Van, 15 km beyond Hoşap, Zernek dam environment, rocky shadows, 1966 m, 27 April 2011, M. Tekşen 2801 & S. Karaman ( AKSU!) ; Güzelsu, 8 km from Hoşap to Günbaşı village , steppe, 1950 m, 15 May 2012, M. Tekşen 2875 & S. Karaman ( AKSU!) ; Güzelsu, 13 km from Hoşap to Günbaşı village , steppe, 1950 m, 15 May 2012, M. Tekşen 2879 & S. Karaman ( AKSU!) ; Güzelsu, 7 km from Hoşap to Gürpınar , steppe, 1971 m, 16 May 2012, M. Tekşen 2882 & S. Karaman ( AKSU!) ; Güzelsu, 5 km from Hoşap to Van, steppe, 1964 m, 14 May 2010, M. Tekşen 2475, 2479 & S. Karaman ( AKSU!) ; Güzelsu, 8.5 km from Hoşap to Van, steppe, 1968 m, 14 May 2010, M. Tekşen 2482 & S. Karaman ( AKSU!) ; Güzelsu, 500 m to Taşdöndüren village , steppe, 2053 m, 14 May 2010, M. Tekşen 2474 & S. Karaman ( AKSU!) ; Güzelsu, 15 km from Hoşap to Van, rocky places, 1990 m, 19 May 2013, M. Tekşen 2919 & S. Karaman ( AKSU!) ; Güzelsu, Güzelsu-Taşdöndüren village , steppe, 1998 m, 14 May 2010, M. Tekşen 2468 & S. Karaman ( AKSU!) ; Gürpınar, Üçgen village-Zernek dam, steppe, 2000 m, 12 May 2007, I. Demir 123 ( VANF!) ; Gürpınar, Akbulut village-Zernek dam, meadows, 2000 m, 22 May 2002, M. Ünal 6792 ( VANF!) ; Gürpınar, above Sapakonak village , meadows, 2547 m, 18 May 2013, M. Tekşen 2904 & S. Karaman ( AKSU!) ; Gürpınar, Işıkpınar-Hacıköy , steppe, 2154 m, 18 May 2013, M. Tekşen 2912 & S. Karaman ( AKSU!) ; Gürpınar, Bölmeçalı- Sapakonak village crossroad, steppe, 2133 m, 13 May 2010, M. Tekşen 2440 & S. Karaman ( AKSU!) ; Gürpınar, Yoldüştü village , meadows, 2242 m, 13 May 2010, M. Tekşen 2447 & S. Karaman ( AKSU!) ; Gürpınar, above Hamurkesen village , steppe, 1965 m, 13 May 2010, M. Tekşen 2456 & S. Karaman ( AKSU!) ; Gürpınar, above Tutmaç village , steppe, 2160 m, 13 May 2010, M. Tekşen 2464 & S. Karaman ( AKSU!) ; Gürpınar, Işıkpınar-Hacıköy , steppe, 2168 m, 13 May 2010, M. Tekşen 2462 & S. Karaman ( AKSU!) ; Erciş- Hica, 5 km beyond Yukarıışıklı village, Koçköprü dam environment, steppe, 1916 m, 29 June 2010, M. Tekşen 2846 & S. Karaman ( AKSU!) ; Erciş, Yukarıışıklı village, Işıklı hill, steppe, rocky places, 1691 m, 8 April 2007, O. Karabacak 5615 ( VANF!) ; Erciş, Altındere stud farm, fields, 1816 m, 12 May 2010, M. Tekşen 2424 & S. Karaman ( AKSU!) ; Erciş, Işbaşı- Hasanabdal village , steppe, 1827 m, 12 May 2010, M. Tekşen 2420 & S. Karaman ; Erciş- Hica, 1–2 km beyond Yukarıışıklı village, Koçköprü dam environment, steppe, 1916 m, 12 May 2010, M. Tekşen 2417 & S. Karaman ( AKSU!) ; Erciş- Hica, Salıklar neighborhood, Yukarıışıklı village , steppe, 1704 m, 12 May 2010, M. Tekşen 2416 & S. Karaman ( AKSU!) ; Erciş, Işbaşı- Hasanabdal village , steppe, 1827 m, 12 May 2010, M. Tekşen 2421 & S. Karaman ( AKSU!) ; Muradiye, above Adaklı village , rocky slopes, 2400 m, 10 May 1998, M. Ünal 2389 ( VANF!) ; Muradiye, Alkasnak village , rock shadows, 1800 m, 31 March 2001, O. Karabacak 1064 ( VANF!) ; Muradiye, above Topuzarpa village, meadows, 1844 m, 12 May 2010, M. Tekşen 2430 ( AKSU!), 2433 & S. Karaman ( GAZI!) ; Muradiye, above Ünseli village, S slopes of Akçadağ , steppe, 1747 m, 12 May 2010, M. Tekşen 2429 & S. Karaman ( AKSU!) ; Özalp, Kalecik village, Ahtadağ , steppe, 2321 m, 21 April 1997, F. Özgökçe 1330 ( VANF!) ; Muradiye, above Yumaklı village , stony places, 1865 m, 12 May 2010, M. Tekşen 2435 & S. Karaman ( AKSU!) ; Gevaş, W of Çadır mountain , steppe, 2300 m, 6 June 1993, Y. Altan 5072 ( GAZI!) ; Erçek, Dereüstü village , rocky places, 1800 m, 30 May 1993, Y. Altan 4812 ( GAZI!) ; Başkale, Çaldıran village , S slopes of Türktepe, steppe, 2050 m, 23 March 2006, D. Avlamaz 568 ( VANF!) .

Flowering time: — April–June.

Habitat: — Rocky and stony places, rock shadows, steppe, 1350–3000 m.

Distribution: — Euro-Siberian element. Turkey (East Anatolia), North Caucasia, Transcaucasia, Central Asia, West Russia, Iran ( Govaerts 2015).

Sect. Didymobulbos (K.Koch) Boiss., Fl. Orient. 5: 204 (1882) ≡ Gagea subsect. Didymobulbos K.Koch, Linnaea 22: 229 (1849) = sect. Fistulosae (Pascher) Davlianidze, Not. Syst. Geogr. Inst. Bot. Thbilissiensis 30: 62 (1973).

Type (lectotype designated by Davlianidze 1973: 62): G. villosa (M.Bieb.) Sweet.

In Turkey, sect. Didymobulbos is the largest section, represented by 13 species. Gagea tenera Pascher (1904: 129) which nomenclatural type was designated by Rix (1984) from Taschkent ( Usbekistan), belongs to sect. Didymobulbos and was cited only for one locality (Van: gorge 8 km West of Hoşap) in Turkey ( Rix 1984). However, Ali & Levichev (2007) excluded this species from Turkey. Despite this, Zarrei et al. (2007) followed Rix (1984) and stated G. tenera as occurring in Turkey. Tison et al. (2013) do not provide certain information about the presence of this species in the country. As a result of our field studies and relevant literature in Van province and surroundings ( Özçelik 1987, Behçet 1989, Öğün & Altan 1992, Altan & Uğurlu 2000, Ünal & Behçet 2007, Bani & Adıgüzel 2008, Pınar & Adıgüzel 2011) we were not able to confirm the occurrence of this species in the country. Moreover, Zernek dam was built at the locality where Rix collected samples. In light of these data, Gagea tenera is here excluded from the flora of Turkey. In addition, as a result of the field studies, examined specimens in the herbaria and the specimens indicated by Rix (1984), the samples which were formerly identified as G. foliosa were indeed referred either to G. dubia , G. ramulosa or G. villosa . In light of these data, we determined that G. foliosa does not occur in Turkey.

Identification key for Gagea sect. Didymobulbos in Turkey:

1. Basal leaf becoming more or less fistulose or always fistulose in cross section................................................................................2

- Basal leaf never fistulose, terete or flattened......................................................................................................................................5

2. Basal leaf with 3 bundles arranged in V; becoming more or less fistulose........................................................................................3

- Basal leaf with 6–11 bundles; always fistulose..................................................................................................................................4

3. Basal leaf distinctly trihedral in outline with a deeply canaliculate adaxial side and strongly convex to almost carinate abaxial side; flowers 1–4(–7) ................................................................................................................................................................ G. lojaconoi

- Basal leaves flatly subtrihedral in outline with a slightly canaliculate adaxial side and a slightly convex abaxial side in basal leaves cross section; flowers 5–12 ........................................................................................................................................... G. antakiensis

4. Basal leaf trihedral in cross section; with 11 bundles; cauline leaves 2–6, opposite, subopposite or verticillate ............. G. fragifera

- Basal leaf round wavy in cross section; with 6 bundles; cauline leaves 2, alternate ....................................................... G. luteoides

5. Basal leaf trihedral to subtrihedral.....................................................................................................................................................6

- Basal leaf flat....................................................................................................................................................................................11

6. Cauline leaves opposite to subopposite..............................................................................................................................................7

- Cauline leaves alternate......................................................................................................................................................................8

7. Cauline leaves twisted; paniculate inflorescence pedicel glabrous................................................................................ G. tenuissima

- Cauline leaves erect; umbellate to subumbellate inflorescence; pedicel hairy ................................................................. G. glacialis

8. Thickened roots present......................................................................................................................................................................9

- Thickened roots absent.....................................................................................................................................................................10

9. Subumbellate inflorescence; pedicel sparsely hairy, with a dense beard of hairs at junction of pedicel and flower ............. G. juliae

- Paniculate inflorescence; pedicel densely hairy, without a dense beard of hairs at junction of pedicel and flower ....... G. bohemica

10. Basal leaf longer than inflorescence; cauline leaves 1; umbellate inflorescence with 1–7 flowers, pedicel densely hairy................. .................................................................................................................................................................................... G. peduncularis

- Basal leaf equalling or shorter than inflorescence; cauline leaves more than 1; corymbose inflorescence with 1–3 flowers, pedicel glabrous or sparsely hairy ................................................................................................................................................. G. bithynica

11. Basal leaves of adult plants with one layer of 5–11 bundles closer to the abaxial epidermis than to the adaxial one; pedicels hairs sometimes short (<1 mm) and straight ................................................................................................................................ G. villosa

- Basal leaves of adult plants with one layer of bundles equally close to the two sides; pedicels hairs, usually> 1 mm and flexuose..........................................................................................................................................................................................................12

12. Basal leaves almost flat, drooping; flowering plants usually with axillary bulbils except at senile stage............................. G. dubia

- Basal leaves canaliculate, rigid, more or less erect or twisted; flowering plants often with axillary bulbils at the first year, then without bulbils .................................................................................................................................................................. G. ramulosa