Takhtajaniantha Nazarova, Biol. Zhurn. Armenii 43: 179. 1990.

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

Takhtajaniantha Nazarova, Biol. Zhurn. Armenii 43: 179. 1990.

= Scorzonera sect. Fibrillosae Nakai, Rep. Inst. Sci. Res. Manchoukuo, ser. 1, 6: 171. 1937. Type: Scorzonera glabra Rupr. [= S. austriaca Willd.]

= Scorzonera subsect. Egregia Kult., Tau-Sagyz Ekol. Osnovy Vvedeniya Ego Kul’t.: 108. 1938.

≡ Scorzonera subg. Egregia (Kult.) Ovcz., Soobshch. Tadzhiksk. Fil. Akad. Nauk SSSR 20: 54. 1950.

≡ Scorzonera sect. Egregia (Kult.) Lipsch., Fl. URSS 29: 58. 1964. Lectotype (designated here): Scorzonera tau-saghyz Lipsch. & G.G.Bosse

= Scorzonera sect. Pusillae Lipsch., Fragm. Monogr. Gen. Scorzonera 2: 59. 1939. Type: Scorzonera pusilla Pall.

= Scorzonera sect. Capito Tzvelev, Rast. Tsentral. Azii 14b: 115. 2009. Type: Scorzonera capito Maxim.

Type species.

Takhtajaniantha pusilla (Pall.) Nazarova

Diagnostic features.

Perennials; caudex often with dark brown fibrous leaf sheath residues and base of the rosulate leaves often lanate; pollen with 6 lacunae.

Description.

Habit, life form, subterranean parts: perennial herbs or subshrublets with a taproot or a tuber.

Leaves: rosulate or crowded in the basal part of the stem, rarely dispersed along the stem, sessile or petiolate, filiform or ovate, apically sometimes hooked ( T. pusilla ), margin flat or undulate, entire or denticulate.

Stem, synflorescence: stem solitary or branched, scape-like or leafy, capitula terminal.

Pollen: echinolophate, tricolporate and each colpus divided into 2 lacunae, with 6 abporal lacunae (sometimes with spineless areas resembling rudimentary lacunae ( Blackmore 1982).

Capitula: glabrous to arachnoid hairy or tomentose, phyllaries in several series, outer phyllaries smaller than inner ones, triangular-ovate, inner phyllaries linear to ovate, receptacle glabrous, flat, with many or rarely few (up to 12: T. pseudodivaricata ) florets; florets yellow (Fig. 16D ) or with dark red veins, usually clearly exeeding the incolucre.

Achenes: 5-15 mm, without carpopodium, straight or slightly curved, often with 10 ribs or rarely roundish, glabrous or hairy, sometimes (as in T. pusilla ) with stout conglomerations; achene wall with parenchyma of two types, with thick- and thin-walled cells, present only as subepidermal continuous layer(s) or only with thick-walled cells, air cavities absent, sclerenchyma with continuous layer with a narrow or wider hunch on either side of the principal rib or, as in T. pseudodivaricata , without hunches, but discontinuous forming 5 bundles (with a gap in the principal ribs), or continuous, its fibres orientated parallel to the fruit axis, tannins absent.

Pappus: 7-28 mm, ice white or dirty white, bristles scabrid in upper part.

Chromosome number: x = 7 (diploids); 2n = 28 in T. pusilla (amphiploid); in T. tau-saghyz polyploid cytoraces 2n = 14, 21, 28, 42 and aneuploids with chromosome numbers 15, 16, 17, 18, 22 and 24, 28 and 42 are known ( Krayevoy 1934; Poddubnaya-Arnoldi et al. 1934; Sosnovets 1960).

Species.

(1) Takhtajaniantha austriaca (Willd.) Zaika, Sukhor. & N.Kilian, comb. nov. (Fig. 16D ) ≡ Scorzonera austriaca Willd., Sp. Pl. 3(3): 1498. 1803 (incl. S. glabra Rupr.). urn:lsid:ipni.org:names:77204033-1

(2) Takhtajaniantha capito (Maxim.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera capito Maxim., Bull. Acad. Imp. Sci. Saint-Pétersb. 32: 491. 1888. urn:lsid:ipni.org:names:77204030-1

(3) Takhtajaniantha crispa (M.Bieb.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera crispa M.Bieb., Fl. Taur.-Caucas. 2: 234. 1808. urn:lsid:ipni.org:names:77204028-1

(4) Takhtajaniantha ikonnikovii (Krasch. & Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera ikonnikovii Krasch. & Lipsch. in Lipschitz, Fragm. Monogr. Gen. Scorzonera 1: 109. 1935. urn:lsid:ipni.org:names:77204024-1

(5) Takhtajaniantha mongolica (Maxim.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera mongolica Maxim., Bull. Acad. Imp. Sci. Saint-Pétersb. 32: 492. 1888. urn:lsid:ipni.org:names:77204020-1

(6) Takhtajaniantha pseudodivaricata (Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera pseudodivaricata Lipsch., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 42: 158. 1933. urn:lsid:ipni.org:names:77204019-1

(7) Takhtajaniantha pusilla (Pall.) Nazarova

(8) Takhtajaniantha subacaulis (Regel) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera austriaca var. subacaulis Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada 6: 323. 1880. urn:lsid:ipni.org:names:77204015-1

(9) Takhtajaniantha tau-saghyz (Lipsch. & G.G.Bosse) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera tau-saghyz Lipsch. & G.G.Bosse, Trudy Nauchno-Issl. Lab. Tresta "Kauchukonos" 4: 27. 1930. urn:lsid:ipni.org:names:77204010-1

(10) Takhtajaniantha veresczaginii (Kamelin & S.V.Smirn.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera veresczaginii Kamelin & S.V.Smirn., Turczaninowia 5(1): 17. 2002. urn:lsid:ipni.org:names:77204008-1

Distribution area.

N Africa: EG. Asia-Temperate: AF; AR; AZ; CN; IL; IR; JO; KG; KP; KR; KZ; MN; RU (Asiatic part); SA; TJ; TM; UZ. Asia-Tropical: PK. Europe: AL; AT; BA; BG; BY; CH; CZ; DE; FR; GR; HR; HU; IT; MD; ME; RO; RS; RU; SI; SK; UA.

The following species may belong to Takhtajaniantha , based on their morphological characters (they were included neither in the carpological nor in molecular investigations):

Scorzonera aniana N.Kilian, S. grubovii Lipsch., S. karataviensis Kult. (possibly a synonym of T. tau-saghyz ), S. manshurica Nakai, S. pamirica C.Shih, S. sinensis (Lipsch. & Krasch.) Nakai

Figures

Figure 16. Images of living plants. A, B Ramaliella tortuosissima A general view B close up view of the capitula (Jordan, Nakeb, 1250 m alt., 8 May 2019). Photograph by O. Fragman-Sapir C Pseudopodospermum papposum (Jordan, Nakeb, 8 May 2019). Photograph by O. Fragman-Sapir D Takhtajaniantha austriaca (Italy, South Tyrol, Schlanders, 6 May 2010). Photograph by R. Hand E Scorzonera humilis (Germany, Ammersee, 15 May 2009). Photograph by C. Niederbichler F Scorzonera meyeri (Russia, Karachaevo-Cherkessiya, Teberda, 14 July 2005). Photograph by A. S. Zernov.