Scorzonera sensu lato (Asteraceae, Cichorieae) - taxonomic reassessment in the light of new molecular phylogenetic and carpological analyses
Author
Zaika, Maxim A.
Author
Kilian, Norbert
Author
Jones, Katy
Author
Krinitsina, Anastasiya A.
Author
Nilova, Maya V.
Author
Speranskaya, Anna S.
Author
Sukhorukov, Alexander P.
text
PhytoKeys
2020
137
1
85
http://dx.doi.org/10.3897/phytokeys.137.46544
journal article
http://dx.doi.org/10.3897/phytokeys.137.46544
1314-2003-137-1
FB2C14EFA257564AA84CD47711BBC273
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 &
Coskunc
., Turk. J. Bot. 39: 77. 2015. Type:
S. zorkunensis
Coskunc
. & 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
(
Suembuel
) Zaika, Sukhor. & N.Kilian, comb. nov. ≡
Scorzonera longiana
Suembuel
, 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 &
Hamzaoglu
,
S. zorkunensis
Coskunc
. & Makbul