Scorzonerinae
Content
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 ‒ native; 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 ‒ native; Korea ‒ native (North Korea ‒ native); Krasnoyarsk ‒ native; Kuwait ‒ native; Lebanon-Syria (Lebanon ‒ native, Syria ‒ native); Manchuria (Heilongjiang ‒ native, Jilin ‒ native, Liaoning ‒ native); Mongolia ‒ native; 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 ‒ native; Tibet ‒ native; Transcaucasus (Abkhaziya ‒ native, Adzhariya ‒ native, Armenia ‒ native, Azerbaijan ‒ native, Georgia ‒ native, Nakhichevan ‒ native); Turkey ‒ native; Turkmenistan ‒ native; Tuva ‒ native; Uzbekistan ‒ native; 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 ‒ native; 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,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
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,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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X. Tremetsberger K., Gemeinholzer B., Zetzsche H. & al., Divergence time estimation in Cichorieae (Asteraceae) using a fossil-calibrated relaxed molecular clock in Organisms Diversity Evol. 13: 1-13. 2012