Scorzonerinae

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Scorzonerinae

Distribution

Africa: Algeria native; Canary Is. native: doubtfully native; Cape Provinces (Eastern Cape Province introduced, Northern Cape Province introduced, Western Cape Province introduced); Egypt native; Ethiopia introduced; Free State introduced; KwaZulu-Natal introduced; Lesotho introduced; Libya native; Madeira native; Morocco native (Morocco native); Northern Provinces (Gauteng introduced, Mpumalanga introduced, North-West Province introduced, Northern Province introduced); Tunisia native Asia-Temperate: Afghanistan nativenative; Altay native; Amur native; Buryatiya native; China North-Central (Beijing introduced, Gansu native, Hebei native, Shaanxi native, Shandong native, Shanxi native); China South-Central (Guizhou native, Hubei native, Sichuan introduced, Yunnan introduced); China Southeast (Anhui native, Henan native, Jiangsu native); Chita native; Cyprus native; East Aegean Is. native; Gulf States (Bahrain native, United Arab Emirates native); Inner Mongolia (Nei Mongol native, Ningxia native); Iran native; Iraq native; Irkutsk native; Kazakhstan native; Khabarovsk native; Kirgizistan nativenative; Korea native (North Korea native); Krasnoyarsk native; Kuwait native; Lebanon-Syria (Lebanon native, Syria native); Manchuria (Heilongjiang native, Jilin native, Liaoning native); Mongolia nativenative; North Caucasus native (Chechnya native, Dagestan native, Kabardino-Balkariya native, Karacheyevo-Cherkessiya native, Krasnodar native, Severo-Osetiya native, Stavropol native); Oman native; Palestine (Israel native, Jordan native); Primorye native; Qinghai native; Sakhalin native; Saudi Arabia native; Sinai native; Tadzhikistan nativenative; Tibet native; Transcaucasus (Abkhaziya native, Adzhariya native, Armenia native, Azerbaijan native, Georgia native, Nakhichevan native); Turkey native; Turkmenistan nativenative; Tuva native; Uzbekistan nativenative; West Siberia native; Xinjiang native; Yakutskiya native; Yemen (North Yemen native, South Yemen native) Asia-Tropical: East Himalaya (Bhutan native); India (Haryana native, Punjab native, Rajasthan native, Uttar Pradesh native); Jawa introduced; Myanmar introduced; Nepal native; Pakistan nativenative; West Himalaya (Himachal Pradesh native, Jammu-Kashmir native) Australasia: New South Wales (Australian Capital Territory introduced, New South Wales introduced); New Zealand North introduced; New Zealand South introduced; Northern Territory introduced; Queensland (Queensland introduced); South Australia introduced; Tasmania introduced; Victoria introduced; Western Australia (Western Australia introduced) Europe: Albania native; Austria native (Austria native, Liechtenstein native); Baleares native; Baltic States (Estonia native, Kaliningrad native, Latvia native, Lithuania native); Belarus native; Belgium native (Belgium native, Luxembourg native); Bulgaria native; Central European Russia native; Corse native; Czechoslovakia native (Czech Republic native, Slovakia native); Denmark native; East European Russia native; Finland native; France native (France native); Germany native; Great Britain native; Greece native; Hungary native; Iceland introduced; Ireland native (Ireland native, Northern Ireland native); Italy native (Italy native); Kriti native; Krym native; Netherlands native; North European Russia native; Northwest European Russia native; Norway native; Poland native; Portugal native; Romania native; Sardegna native; Sicily native (Malta native, Sicily native); South European Russia native; Spain native (Andorra native, Gibraltar native, Spain native); Sweden native; Switzerland native; Turkey-in-Europe native; Ukraine (Moldova native, Ukraine native); Yugoslavia (Bosnia-Herzegovina native, Croatia native, Macedonia native, Montenegro native, Serbia native, Slovenia native) Northern America: Alberta introduced; Arizona native; Arkansas introduced; British Columbia introduced; California introduced; Colorado introduced; Connecticut introduced; Delaware introduced; District of Columbia introduced; Georgia, U.S.A. native; Idaho native; Illinois introduced; Indiana introduced; Iowa introduced; Kansas introduced; Kentucky introduced; Labrador introduced; Louisiana introduced; Maine introduced; Manitoba introduced; Maryland introduced; Massachusetts introduced; Mexico Northwest (Baja California introduced); Michigan introduced; Minnesota introduced; Missouri introduced; Montana introduced; Nebraska introduced; Nevada introduced; New Brunswick introduced; New Hampshire introduced; New Jersey introduced; New Mexico introduced; New York introduced; Newfoundland (Newfoundland introduced); North Carolina introduced; North Dakota introduced; Northwest Territories introduced; Nova Scotia introduced; Ohio introduced; Oklahoma introduced; Ontario introduced; Oregon introduced; Pennsylvania introduced; Prince Edward I. introduced; Québec introduced; Rhode I. introduced; Saskatchewan introduced; South Dakota introduced; Tennessee introduced; Texas introduced; Utah introduced; Vermont introduced; Virginia introduced; Washington native; West Virginia introduced; Wisconsin introduced; Wyoming native; Yukon introduced Southern America: Argentina Northeast (Buenos Aires introduced, Córdoba introduced, La Pampa introduced); Argentina Northwest (Jujuy introduced, Mendoza introduced, San Juan introduced, San Luis introduced); Argentina South (Chubut introduced, Neuquén introduced, Rio Negro introduced, Santa Cruz introduced, Tierra del Fuego (Argentina) introduced); Chile Central (Biobío introduced, Coquimbo introduced, O'Higgins introduced, Santiago introduced, Valparaíso introduced); Chile South (Aisén introduced); Dominican Republic introduced; Haiti (Haiti introduced); Puerto Rico introduced

Systematics

The subtribe Scorzonerinae forms a well-supported clade in its traditional morphological characterisation and circumscription (Stebbins 1953, but lacking Koelpinia; Blackmore 1981; Bremer 1994; Lack 2006) in all phylogenetic analyses of the Cichorieae based on morphological (Bremer 1994) and molecular data (Whitton et al. 1995; Mavrodiev & al. 2004; Kilian & al. 2009; Tremetsberger et al. 2013).

The Scorzonerinae are characterised by predominantly linear-lanceolate and parallel-veined leaves, an indumentum being soft or absent, uni- to multiseriate involucral bracts, a softly fimbriate plumose pappus and an echinolophate pollen type of its own having colpori with only two (in all other members of the tribe with three) lacunae (Blackmore 1986). If the pappus is absent as in Koelpinia, the characteristic pollen type of this alliance still allows unequivocal placement (Blackmore 1981).

The genus Avellara, with its single species A. fistulosa, a rare endemic of the Iberian Peninsula formerly placed in Scorzonera, has to be excluded from the Scorzonerinae for morphological, palynological, cytological (Blanca & Díaz 1985) and molecular phylogenetic evidence (Fernández-Mazuecos & al. 2016). It apparently represents an isolated, early diverging lineage of the large Chondrillinae-Crepidinae-Hyoseridinae-Hypochaeridinae-Lactucinae clade (Kilian & al. 2009, Tremetsberger & al. 2013) of the Cichorieae with the closest affinities probably to the Hypochaeridinae. For the time being, this genus is therefore treated as a member of that subtribe.

Generic delimitation within the subtribe has been controversial, mainly regarding the circumscription of Scorzonera, the largest and name-giving genus of the subtribe. Zaika & al. (2020: 3) give a brief summary of the taxonomic history: "Differences in leaf or achene shape and pubescence were soon after the establishment of the genus used to separate some taxa into the genera Podospermum (Candolle in Lamarck & Candolle 1805), Gelasia (Cassini 1818) and Lasiospora (Cassini in Cuvier 1822), but only Scorzonera and Podospermum were accepted in the first comprehensive works on Asteraceae systematics by Lessing (1832) and Candolle (1838). Subsequent authors accepted Scorzonera again in a broader sense, with more or less elaborated infrageneric classification (Bentham in Bentham and Hooker 1873; Boissier 1875; Hoffmann 1890–1894; Lipschitz 1935, 1939, 1964). Three major entities were mostly accepted in the 20th century: Scorzonera s.str., Podospermum and Pseudopodospermum, either at subgeneric rank (Lipschitz in Bobrov & Tzvelev 1964; Chamberlain 1975; Chater in Tutin & al. 1976; Rechinger 1977) or, more rarely, at generic rank (Kuthatheladze 1978)."

Molecular phylogenetic analyses of the subtribe by Mavrodiev & al. (2004), Winfield & al. (2006) clearly revealed the polyphyly of Scorzonera in its wide sense. Zaika & al. (2020) confirmed the polyphyly of Scorzonera and presented a re-evaluation of Scorzonera, based on carpological (including anatomical) and molecular phylogenetic analyses of a comprehensive sampling, spanning, for the first time, the entire subtribe and all major groups.

Morphology, even including fruit anatomy, does not, however, very well reflect the structure of the subtribe as revealed through molecular phylogenetics. Actually, most of the lineages resolved are difficult to characterise by morphology and even more distant clades of the Scorzonerinae are often not well distinguished morphologically (Zaika & al. 2020).The ITS phylogeny by Zaika & al. (2020) shows three principal options for a revised classification based on monophyletic generic concepts. The first is to separate generically the Gelasia clade and retain the remainder of Scorzonera s.l. with the Pterachaenia, Koelpinia, Takhtajaniantha, Pseudopodospermum and Podospermum clades included in it. However, the actual inconvenience and challenge is morphologically distinghuishing Gelasia as separate from Scorzonera. This burden is inevitable and shared by all three available options. The second option is to recognise Scorzonera in a sense to include also the Takhtajaniantha, Pseudopodospermum and Podospermum clades. The third option, proposed by Zaika & al. (2020) and equally resolved in their nrITS and plastid DNA tree, is to recognise Scorzonera in a narrow sense to include only the sister clades Scorzonera s.str.and Podospermum.

The revised generic classification of the Scorzonerinae by Zaika & al. (2020), which is followed here, recognises seven genera in accordance with the phylogenetic lineages resolved: Gelasia, Lipschitzia (a new genus for the monospecifc Scorzonera divaricata clade), Pseudopodospermum, Pterachaenia, Ramaliella (a new genus accommodating the S. polyclada clade), Scorzonera and Takhtajaniantha.A,AA,AB,B,C,D,E,F,G,H,I,J,K,L,M,N,O,P,Q,R,S,T,U,V,W,X,Y,Z

Bibliography

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