Gelasia Cass., Bull. Sci. Soc. Philom. Paris 1818: 33. 1818.

Zaika, Maxim A., Kilian, Norbert, Jones, Katy, Krinitsina, Anastasiya A., Nilova, Maya V., Speranskaya, Anna S. & Sukhorukov, Alexander P., 2020, Scorzonera sensu lato (Asteraceae, Cichorieae) - taxonomic reassessment in the light of new molecular phylogenetic and carpological analyses, PhytoKeys 137, pp. 1-85 : 1

Gelasia Cass., Bull. Sci. Soc. Philom. Paris 1818: 33. 1818.

≡ Scorzonera sect. Gelasia (Cass.) DC., Prodr. 7(1): 123. 1838. Type: Scorzonera villosa Scop. ≡ Gelasia villosa (Scop.) Cass.

= Lasiospora Cass. in Cuvier, Dict. Sci. Nat. 25: 306. 1822.

≡ Scorzonera sect. Lasiospora (Cass.) Less., Syn. Gen. Compos.: 134. 1832.

Lasiospora ≡ Scorzonera subg. Lasiospora (Cass.) Peterm., Deutschl. Fl.: 334. 1846-1849. Lectotype ( Tzvelev 1989: 45): Lasiospora hirsuta (Gouan) Cass.

= Scorzonera subsect. Pulvinares Boiss., Fl. Orient. 3: 756. 1875.

≡ Scorzonera sect. Pulvinares (Boiss.) Lipsch., Fragm. Monogr. Gen. Scorzonera 1: 24. 1935. Lectotype (designated here): Scorzonera seidlitzii Boiss.

= Scorzonera subsect. Infrarosulares Hand.-Mazz., Ann. K. K. Naturhist. Hofmus. 27: 455. 1913.

≡ Scorzonera sect. Infrarosulares (Hand.-Mazz.) Lipsch., Fragm. Monogr. Gen. Scorzonera 1: 10. 1935. Lectotype (designated here): Scorzonera acantholimon Hand.-Mazz.

= Scorzonera sect. Nervosae Lipsch., Fragm. Monogr. Gen. Scorzonera 1: 126. 1935. Type: Scorzonera latifolia (Fisch. & C.A.Mey.) DC.

= Scorzonera sect. Trachyactis Rech.f., Oesterr. Bot. Z. 84: 169. 1935. Lectotype (designated here): Scorzonera cretica Willd. Note: This section was described, based on the entirely scabrid pappus, a remarkable feature in Gelasia and the subtribe. Three species were reported to form this section: S. dependens Rech.f. (= S. cretica Willd.), S. araneosa Sm. and S. eximia Rech.f. The first species (as S. cretica ) was nested in our nrITS phylogeny in the Gelasia clade. Two species, S. cretica and S. araneosa , were included in our carpological analysis and match the carpology of Gelasia .

= Scorzonera sect. Tuberosae Lipsch., Fragm. Monogr. Gen. Scorzonera 2: 39. 1939. Type (Art. 10.8, see Turland et al. 2018): S. tuberosa Pall.

= Scorzonera sect. Vierhapperia Lipsch., Fragm. Monogr. Gen. Scorzonera 2: 88. 1939. Lectotype (designated here): Scorzonera ensifolia M.Bieb.

= Scorzonera sect. Anatolia Makbul & Coşkunç., Turk. J. Bot. 39: 77. 2015. Type: S. zorkunensis Coskunç. & Makbul

Diagnostic features.

Perennials; leaves entire; involucre often pubescent; achenes often densely lanate, rarely glabrous, usually smooth rarely with emergence; pappus often rigid, plumose in lower portion and scabrid in upper portion or rarely almost completely scabrid; sclerenchyma forming a sheath, with an invagination on either side of the principal rib.

Description.

Habit, life form, subterranean parts: perennial herbs of various habits, rarely subshrubs (only G. acantholimon ), with a taproot, more rarely with globose or cylindrical tuber.

Leaves: rosulate or cauline, sessile or petiolate, linear to ovate, often hairy (frequently very densely so, fleecy, lanate) or glabrous, margin entire and flat or undulate.

Stem, synflorescence: scape-like, small or leafy, tall and branched, often pubescent, more rarely glabrous (usually like those of the leaves), synflorescence racemiform, spiciform or corymbiform or capitula single.

Capitula: involucre usually pubescent, sometimes glabrescent at fruiting or rarely glabrous, phyllaries in several series, all lanceolate or triangular, outer phyllaries <3-4 times smaller than the inner ones, receptacle glabrous or hairy, capitula of many yellow florets (Fig. 15C ) or, rarely, only 4-6 ( G. acantholimon ), well exceeding the involucre.

Pollen: echinolophate, tricolporate and each colpus divided into 2 lacunae; with 18 (6 abporal, 6 equatorial and 6 interporal) or, more rarely, 24 (in addition with 6 polar: G. lanata , G. pygmaea ) lacunae ( Blackmore 1982; Pinar et al. 2016), extensive echinate polar areas present.

Achenes: 4-12 mm long, without carpopodium, often densely covered with long woolly indumentum, more rarely glabrous, straight or slightly curved, 5-ribbed or roundish, smooth or rarely with stout conglomerations ( G. caespitosa , G. pygmaea and G. villosa ), parenchyma usually insular above invaginations of the sclerenchyma and below the principal ribs or only insular above invaginations of sclerenchyma, air cavities absent, sclerenchymatous layers continuous or discontinuous in the main ribs, forming a sheath, with an invagination on either side of the principal rib, fibres orientated parallel to the fruit axis, tannins absent.

Pappus: 5-24 mm; bristles plumose in lower portion and scabrid in upper portion or more rarely, almost completely scabrid; dirty white, yellow or fulvous.

Chromosome number.

x = 6 (rarely 7), diploids or tetraploid.

Species.

(1) Gelasia acantholimon (Hand.-Mazz.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera acantholimon Hand.-Mazz. in Ann. K. K. Naturhist. Hofmus. 27: 455. 1913. urn:lsid:ipni.org:names:77204005-1

(2) Gelasia albicans (Coss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera albicans Coss., Notes Crit.: 11. 1851. urn:lsid:ipni.org:names:77204006-1

(3) Gelasia araneosa (Sm.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera araneosa Sm. in Sibthorp, Fl. Graec. Prodr. 2(1): 123. 1813. urn:lsid:ipni.org:names:77204007-1

(4) Gelasia aucheriana (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera aucheriana DC., Prodr. 7(1): 125. 1838. urn:lsid:ipni.org:names:77204009-1

(5) Gelasia biebersteinii (Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera biebersteinii Lipsch., Fragm. Monogr. Gen. Scorzonera 2: 95. 1939. urn:lsid:ipni.org:names:77204011-1

(6) Gelasia caespitosa (Pomel) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera caespitosa Pomel, Nouv. Mat. Fl. Atl.: 266. 1875. urn:lsid:ipni.org:names:77204012-1

(7) Gelasia callosa (Moris) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera callosa Moris, Stirp. Sard. Elench. 1: 29. 1827. urn:lsid:ipni.org:names:77204014-1

(8) Gelasia cinerea (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera cinerea Boiss., Diagn. Pl. Orient., ser. 1, 11: 44. 1849. urn:lsid:ipni.org:names:77204016-1

(9) Gelasia circumflexa (Krasch. & Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera circumflexa Krasch. & Lipsch., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., ser. 2, 42. 148. 1934. urn:lsid:ipni.org:names:77204017-1

(10) Gelasia cretica (Willd.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera cretica Willd., Sp. Pl. 3: 1504. 1803. urn:lsid:ipni.org:names:77204018-1

(11) Gelasia doriae (Degen & Bald.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera doriae Degen & Bald., Oesterr. Bot. Z. 46: 417. 1896. urn:lsid:ipni.org:names:77204021-1

(12) Gelasia dzhawakhetica (Sosn. ex Grossh.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera dzhawakhetica Sosn. ex Grossh., Fl. Kavk. 4: 236. 1934. urn:lsid:ipni.org:names:77204022-1

(13) Gelasia ensifolia (M.Bieb.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera ensifolia M.Bieb., Fl. Taur.-Caucas. 2: 235. 1808. urn:lsid:ipni.org:names:77204023-1

(14) Gelasia eriophora (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera eriophora DC., Prodr. 7(1): 125. 1838. urn:lsid:ipni.org:names:77204025-1

(15) Gelasia filifolia (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera filifolia Boiss., Fl. Orient. 3: 774. 1875. urn:lsid:ipni.org:names:77204026-1

(16) Gelasia hirsuta (Gouan) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Tragopogon hirsutus Gouan, Fl. Monsp.: 342. 1764. urn:lsid:ipni.org:names:77204027-1

(17) Gelasia ketzkhovelii (Grossh.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera ketzkhovelii Sosn., Trudy Tbilissk. Bot. Inst. 2: 219. 1938. urn:lsid:ipni.org:names:77204029-1

(18) Gelasia kotschyi (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera kotschyi Boiss., Fl. Orient. 3: 780. 1875. urn:lsid:ipni.org:names:77204031-1

(19) Gelasia lanata (L.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Leontodon lanatus L., Cent. Pl. 1: 26. 1755. urn:lsid:ipni.org:names:77204032-1

(20) Gelasia lasiocarpa (Chamberlain) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera lasiocarpa Chamberlain, Notes Roy. Bot. Gard. Edinburgh 33: 255. 1974. urn:lsid:ipni.org:names:77204034-1

(21) Gelasia latifolia (Fisch. & C.A.Mey.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Lasiospora latifolia Fisch. & C.A.Mey., Index Sem. Hort. Petrop. 1: 30. 1835. urn:lsid:ipni.org:names:77204035-1

(22) Gelasia litwinowii (Krasch. & Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera litwinowii Krasch. & Lipsch., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., ser. 2, 43: 153. 1934. urn:lsid:ipni.org:names:77204036-1

(23) Gelasia longiana ( Sümbül) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera longiana Sümbül, Edinburgh J. Bot. 48(1): 35. 1991. urn:lsid:ipni.org:names:77204039-1

(24) Gelasia mackmeliana (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera mackmeliana Boiss., Diagn. Pl. Orient., ser. 1, 11: 44. 1849. urn:lsid:ipni.org:names:77204041-1

(25) Gelasia mirabilis (Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera mirabilis Lipsch., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 22: 288. 1963. urn:lsid:ipni.org:names:77204043-1

(26) Gelasia pisidica (Hub.-Mor.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera pisidica Hub.-Mor., Bauhinia 7(3): 179. 1982. urn:lsid:ipni.org:names:77204044-1

(27) Gelasia psychrophila (Boiss. & Hausskn.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera psychrophila Boiss. & Hausskn. in Boissier, Fl. Orient. 3: 777. 1875. urn:lsid:ipni.org:names:77204045-1

(28) Gelasia pygmaea (Sm.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera pygmaea Sm., Fl. Graec. Prodr. 2(1): 123. 1813. urn:lsid:ipni.org:names:77204046-1

(29) Gelasia ramosissima (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera ramosissima DC., Prodr. 7(1): 125. 1838. urn:lsid:ipni.org:names:77204047-1

(30) Gelasia rigida (Aucher ex DC.) Zaika, Sukhor. & N.Kilian, comb. nov. (Fig. 15 D–F ) ≡ Scorzonera rigida Aucher ex DC., Prodr. 7(1): 123. 1838. urn:lsid:ipni.org:names:77204048-1

(31) Gelasia sandrasica (Hartvig & Strid) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera sandrasica Hartvig & Strid, Bot. Jahrb. Syst. 108(2-3): 311. 1987. urn:lsid:ipni.org:names:77204050-1

(32) Gelasia seidlitzii (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera seidlitzii Boiss., Fl. Orient. 3: 775. 1875. urn:lsid:ipni.org:names:77204051-1

(33) Gelasia sericea (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera sericea DC., Prodr. 7(1): 123. 1838. urn:lsid:ipni.org:names:77204053-1

(34) Gelasia tomentosa (L.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera tomentosa L., Sp. Pl., ed. 2, 2: 1112. 1763. urn:lsid:ipni.org:names:77204054-1

(35) Gelasia tuberosa (Pall.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera tuberosa Pall., Reise Russ. Reich. 3: 757. 1776. urn:lsid:ipni.org:names:77204057-1

(36) Gelasia ulrichii (Parolly & N.Kilian) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera ulrichii Parolly & N.Kilian, Willdenowia 32: 198. 2002. urn:lsid:ipni.org:names:77204058-1

(37) Gelasia villosa (Scop.) Cass.

Distribution area.

N Africa: DZ; EG; LY; MA; TN. Europe: AL; BA; CY; ES; FR; GR; HR; IT; MD; ME; RS; RU; SI; UA. Asia-Temperate: AF; AM; AZ; CH; GE; IL; IQ; IR; JO; KG; KZ; LB; RU (Asiatic part); SY; TJ; TM; UZ. Asia-Tropical: PK.

The following species may also belong to Gelasia , based on their morphological characters (neither included in the carpological nor in molecular analyses): Scorzonera amasiana Hausskn. & Bornm., S. boissieri Lipsch., S. bungei Krasch. & Lipsch., S. charadzae Papava, S. gageoides Boiss., S. ispahanica Boiss. ≡ Lasiospora ispahanica (Boiss.) Walp., S. joharchii S.R.Safavi, S. karabelensis Parolly & N.Kilian, S. kozlowskyi Sosn. ex Grossh., S. lipskyi Lipsch., S. persica Boiss. & Buhse, S. pulchra Lomak., S. safievii Grossh., S. sahnea Parsa, S. scopariiformis Lipsch., S. sublanata Lipsch., S. sericeolanata (Bunge) Krasch. & Lipsch., S. veratrifolia Fenzl ≡ Lasiospora veratrifolia (Fenzl) Walp., S. wendelboi Rech.f., S. woronowii Krasch., S. xylobasis Rech.f., S. yildirimlii A.Duran & Hamzaoğlu, S. zorkunensis Coşkunç. & Makbul

Figures

Figure 15. Images of living plants. A, B - Tourneuxia variifolia (Morocco, near Mrhimina). Photographs by A. Garcin (www. teline. fr) C - Gelasia villosa (cult. in BG Berlin, 13 Jun 2010). Photograph by N. Kilian D, E, F - Gelasia rigida (Jordan, Nakeb, 1250 m alt., 8 May 2019). Photograph by O. Fragman-Sapir. D general view E capitula in flowering F capitula in fruiting, a part of achenes had fallen off G Geropogon hybridus (Greece, Lefkas, 15 May 2016). Photograph by E. Willing H Koelpinia linearis (Qatar, Khashem Al Nekhsh area, 18 Mar 2016). Photograph by A. Sergeev.