Pseudopodospermum (Lipsch. & Krasch.) Kuth., Kavkaz. Predstav. Scorzonerinae: 85. 1978.

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

Pseudopodospermum (Lipsch. & Krasch.) Kuth., Kavkaz. Predstav. Scorzonerinae: 85. 1978.

≡ Scorzonera sect. Pseudopodospermum Lipsch. & Krasch. in Lipschitz, Fragm. Monogr. Gen. Scorzonera 1: 70. 1935.

≡ Scorzonera subg. Pseudopodospermum (Lipsch. & Krasch.) Lipsch. in Bobrov & Tzvelev, Fl. URSS 29: 48. 1964. Lectotype (designated by Lipschitz 1964: 49): Scorzonera mollis M.Bieb.

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

≡ Scorzonera sect. Foliosae (Boiss.) Lipsch. in Bobrov & Tzvelev, Fl. URSS 29: 721. 1964. Lectotype (designated by Lipschitz 1964: 721): S. stricta Hornem.

= Scorzonera sect. Incisae Lipsch., Fragm. Monogr. Gen. Scorzonera 1: 91. 1935.

≡ Podospermum sect. Incisae (Lipsch.) Kuth., Kavkaz. Predstav. Scorzonerinae : 99. 1978. Type (Art. 10.8, see Turland et al. 2018): Scorzonera incisa DC.

= Scorzonera sect. Papposae Lipsch. & Krasch. in Lipsch., Fragm. Monogr. Gen. Scorzonera 1: 45. 1935. Type (Art. 10.8, see Turland et al. 2018): Scorzonera papposa DC.

= Scorzonera ser. Brevicaulis Lipsch., Fragm. Monogr. Gen. Scorzonera 1: 92. 1935. Type (Art. 10.8, see Turland et al. 2018): Scorzonera brevicaulis Vahl

= Scorzonera subsect. Hissarica Lipsch., Fragm. Monogr. Gen. Scorzonera 2: 63. 1939.

≡ Scorzonera sect. Hissarica Lipsch. in Bobrov & Tzvelev, Fl. URSS 29: 97. 1964. Type (Art. 10.8, see Turland et al. 2018): Scorzonera hissarica C.Winkl.

Diagnostic features.

Perennial herbs often with a tuber; leaves entire, often with undulate margin or pinnatifid; capitula solitary or few; pollen with 20 lacunae; achenes usually papillate, with stout conglomerations or verrucose ribs.

Description.

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

Leaves: rosulate only or cauline, lower leaves usually petiolate, upper leaves sessile and often (sub) amplexicaule, their shape diverse (linear-lanceolate to broadly ovate), margin entire and then often undulate, or dentate to pinnatisect.

Stem, synflorescence: stem solitary or with a few branches, sometimes leafless and scapose, glabrous to lanate; capitula terminal.

Capitula: involucre glabrous to lanate, phyllaries in several series, outer phyllaries triangular or ovate, obtuse, inner phyllaries oblong or lanceolate, much longer than outer phyllaries, receptacle naked, flat, florets more than 12, exceeding the involucre up to 1.5 times, yellow or purple (Fig. 16C ), sometimes bicoloured, with yellow ligulae and dark purple mouth.

Pollen: echinolophate, tricolporate and each colpus divided into 2 lacunae, with 20 (6 abporal, 6 equatorial, 6 interporal, 2 polar) lacunae ( Blackmore 1982; Pinar et al. 2016).

Achenes: 8-28 mm, straight or slightly curved, with or without carpopodium, usually with five, rarely with ten ribs or without, surface papillate and/or usually with stout conglomerations or the outer strongly rugose or tuberculate-squamate; achene wall with parenchyma subepidermal, continuous (or very rarely sometimes irregularly discontinuous), often reduced to one- to several layers consisting of thin- and/or thick-walled cells, sometimes with subepidermal mechanical parenchyma, air cavities absent or present, sclerenchyma discontinuous with a gap in the principal ribs or continuous forming a sheath, equal in thickness or with a narrow or wider hunch on either side of the principal rib, sclerenchymatous fibres orientated parallel to the achene axis, tannins absent or present in the cell protoplasts.

Pappus: 6-28 mm, bristles equal or unequal (5-10 bristles often longer than the rest), plumose in lower and scabrid in upper part, dirty white or white, yellowish, blackish or rarely fulvous.

Chromosome number.

x = 7, diploids, tetraploids or hexaploids.

Species.

(1) Pseudopodospermum boeticum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera montana var. boetica Boiss. in Candolle, Prodr. 7(1): 121. 1838 ≡ Scorzonera boetica (Boiss.) Boiss., Voy. Bot. Espagne 2: 382. 1841. urn:lsid:ipni.org:names:77204074-1

(2) Pseudopodospermum brevicaule (Vahl) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera brevicaulis Vahl, Symb. Bot. 2: 88. 1791. urn:lsid:ipni.org:names:77204076-1

(3) Pseudopodospermum calyculatum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera calyculata Boiss., Diagn. Pl. Orient., ser. 1, 11: 42. 1849. urn:lsid:ipni.org:names:77204078-1

(4) Pseudopodospermum chantavicum (Pavlov) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera chantavica Pavlov, Vestn. Akad. Nauk Kazakhsk. S.S.R. 8: 27. 1950. urn:lsid:ipni.org:names:77204079-1

(5) Pseudopodospermum crispatulum (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera hispanica var. crispatula DC., Prodr. 7(1): 121. 1838 ≡ Scorzonera crispatula (DC.) Boiss., Voy. Bot. Espagne 2: 741. 1845. urn:lsid:ipni.org:names:77204082-1

(6) Pseudopodospermum crocifolium (Sm.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera crocifolia Sm. in Sibthorp, Fl. Graec. Prodr. 2(1): 123. 1813. urn:lsid:ipni.org:names:77204084-1

(7) Pseudopodospermum davisii (Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera davisii Lipsch., Bot. Mater. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 22: 291. 1963. urn:lsid:ipni.org:names:77204085-1

(8) Pseudopodospermum elatum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera elata Boiss., Diagn. Pl. Orient., ser. 1, 4: 25. 1844. urn:lsid:ipni.org:names:77204086-1

(9) Pseudopodospermum gracile (Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera gracilis Lipsch. in Bobrov & Tzvelev, Fl. URSS 29: 721. 1964. urn:lsid:ipni.org:names:77204088-1

(10) Pseudopodospermum hispanicum (L.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera hispanica L., Sp. Pl. 2: 791. 1753. urn:lsid:ipni.org:names:77204090-1

(11) Pseudopodospermum hissaricum (C.Winkl.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera hissarica C.Winkl., Trudy Imp. S.-Peterburgsk. Bot. Sada 11: 172. 1889. urn:lsid:ipni.org:names:77204092-1

(12) Pseudopodospermum idaeum (Gand.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Podospermum idaeum Gand., Bull. Soc. Bot. France 62: 155. 1916. urn:lsid:ipni.org:names:77204093-1

(13) Pseudopodospermum inaequiscapum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera inaequiscapa Boiss., Fl. Orient. 3: 762. 1875. urn:lsid:ipni.org:names:77204091-1

(14) Pseudopodospermum incisum (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera incisa DC., Prodr. 7(1): 119. 1838. urn:lsid:ipni.org:names:77204089-1

(15) Pseudopodospermum inconspicuum (Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera inconspicua Lipsch., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., n.s. 42: 128. 1933. urn:lsid:ipni.org:names:77204087-1

(16) Pseudopodospermum lamellatum (Krasch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera lamellata Krasch., Trudy Bot. Inst. Akad. Nauk S.S.S.R., ser. 1, Fl. Sist. Vyssh. Rast. 1: 180. 1933. urn:lsid:ipni.org:names:77204083-1

(17) Pseudopodospermum leptophyllum (DC.) Kuth.

(18) Pseudopodospermum libanoticum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera libanotica Boiss., Diagn. Pl. Orient., ser. 1, 11: 43. 1849. urn:lsid:ipni.org:names:77204081-1

(19) Pseudopodospermum molle (M.Bieb.) Kuth.

(20) Pseudopodospermum mucidum (Rech.f., Aellen & Esfand.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera mucida Rech.f., Aellen & Esfand., Oesterr. Bot. Z. 97: 264. 1950. urn:lsid:ipni.org:names:77204077-1

(21) Pseudopodospermum ovatum (Trautv.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera ovata Trautv., Trudy Imp. S.-Peterburgsk. Bot. Sada 1: 275. 1871. urn:lsid:ipni.org:names:77204075-1

(22) Pseudopodospermum pachycephalum (Podlech & Rech.f.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera pachycephala Podlech & Rech.f. in Rechinger, Fl. Iranica 122: 36. 1977. urn:lsid:ipni.org:names:77204073-1

(23) Pseudopodospermum papposum (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera papposa DC., Prodr. 7(1): 119. 1838. urn:lsid:ipni.org:names:77204072-1

(24) Pseudopodospermum phaeopappum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Podospermum phaeopappum Boiss., Diagn. Pl. Orient., ser. 1, 7: 5. 1846. urn:lsid:ipni.org:names:77204070-1

(25) Pseudopodospermum pubescens (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera pubescens DC., Prodr. 7(1): 122. 1838. urn:lsid:ipni.org:names:77204067-1

(26) Pseudopodospermum raddeanum (C.Winkl.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera raddeana C.Winkl., Trudy Imp. S.-Peterburgsk. Bot. Sada 11: 150. 1889. urn:lsid:ipni.org:names:77204062-1

(27) Pseudopodospermum reverchonii (O.Debeaux ex Hervier) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera reverchonii O.Debeaux ex Hervier, Bull. Acad. Int. Géogr. Bot., sér. 3, 15(187-188): 107. 1905. urn:lsid:ipni.org:names:77204056-1

(28) Pseudopodospermum semicanum (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera semicana DC., Prodr. 7(1): 119. 1838. urn:lsid:ipni.org:names:77204055-1

(29) Pseudopodospermum strictum (Hornem.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera stricta Hornem., Hort. Bot. Hafn.: 2: 750. 1815. urn:lsid:ipni.org:names:77204052-1

(30) Pseudopodospermum suberosum (K.Koch) Kuth., Kavkaz. Predstav. Scorzonerinae : 94. 1978

(31) Pseudopodospermum syriacum (Boiss. & C.I.Blanche) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera syriaca Boiss. & C.I.Blanche, Diagn. Pl. Orient., ser. 2, 3: 93. 1856. urn:lsid:ipni.org:names:77204049-1

(32) Pseudopodospermum szowitzii (DC.) Kuth., Kavkaz. Predstav. Scorzonerinae : 92. 1978.

(33) Pseudopodospermum troodeum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera troodea Boiss., Fl. Orient., suppl.: 320. 1888. urn:lsid:ipni.org:names:77204042-1

(34) Pseudopodospermum turcomanicum (Krasch. & Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera turcomanica Krasch. & Lipsch. in Lipschitz, Fragm. Monogr. Gen. Scorzonera 1: 80. 1935. urn:lsid:ipni.org:names:77204040-1

(35) Pseudopodospermum undulatum (Vahl) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera undulata Vahl, Symb. Bot. 2: 86. 1791. urn:lsid:ipni.org:names:77204038-1

(36) Pseudopodospermum violaceum (Chamberlain) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonera violacea Chamberlain, Notes Roy. Bot. Gard. Edinburgh 33(2): 256. 1974. urn:lsid:ipni.org:names:77204037-1

Distribution area.

N Africa: DZ; LY; MA; TN. Europe: AL; AT; BA; BE; BG; CH; CY; CZ; DE; ES; FR; GR; HR; HU; IT; MD; ME; MK; PL; PT; RO; RS; RU; SK; SI; UA. Asia-Temperate: AF; AM; AZ; EG; GE; IL; IQ; IR; JO; KW; KZ; LB; OM; SY; SA; TJ; TK; TR; UZ. Asia-Tropical: PK. Pseudopodospermum hispanicum as an edible crop was introduced in further regions.

The following species may belong to Pseudopodospermum , based on their morphological characters (they were included neither in the carpological nor in molecular investigations): Scorzonera acuminata Boiss., S. aragatzii Kuth. ≡ Pseudopodospermum aragatzii (Kuth.) Kuth., S. coriacea A.Duran & Aksoy, S. crassifolia Krasch. & Lipsch., S. drarii Täckh., S. ferganica Krasch., S. gorovanica Nazarova, S. helodes Rech.f., S. karkasensis S.R.Safavi, S. lacera Boiss. & Balansa, S. limnophila Boiss., S. mariovoensis Micevski, S. mucida Rech.f. et al., S. pacis Güzel et al., S. nivalis Boiss. & Hausskn., S. rawii Rech.f. & Guest, S. scyria M.A.Gust. & Snogerup, S. serpentinica Rech.f., S. stenocephala Boiss., S. tadshikorum Krasch. & Lipsch., S. tunicata Rech.f. & Köie, S. tuzgoluensis A.Duran et al.

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.