Scorzonerinae

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Systematics

The subtribe Scorzonerinae forms a well-supported clade in all phylogenetic analyses of the Cichorieae based on morphological (Bremer 1994) and molecular data (Mavrodiev & al. 2004; Gemeinholzer & al. in Kilian & al. 2009). The molecular analyses with high statistical support of the monophyly confirm its recognition as subtribe Scorzonerinae in its traditional morphological characterisation and circumscription (Stebbins 1953, but lacking Koelpinia; Blackmore 1981; Bremer 1994; Lack 2006).

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 distinct types of echinolophate pollen (with equatorial ridges replaced by a lacuna, the aperture being divided into two lacunae, and with a characteristic exine stratification). 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 and cytological evidence as was shown by Blanca & Díaz (1985). It apparently represents an isolated, early diverging lineage of the large Chondrillinae-Crepidinae-Hyoserdinae-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 and the recognition of the segregates Epilasia, Podospermum, Pterachaenia, Takhtajaniantha and Tourneuxia. A recent molecular phylogeny of the subtribe by Mavrodiev & al. (2004) confirmed the polyphyly of Scorzonera and provided support for recognition of the aforementioned segregates as well as for the separation of Geropogon from Tragopogon. Further molecular analyses of the subtribe (Gemeinholzer & al., in prep.) including more sequences from the core of Scorzonera revealed paraphyletic groupings even within Scorzonera s.str. of Mavrodiev & al. (2004). However, due to the yet incomplete taxon sampling a revised taxonomic treatment of Scorzonera is not yet available, as this is still subject to ongoing studies (Gemeinholzer & al., in prep.).

References


Blackmore S. 1981: Palynology and intergeneric relationships in subtribe Hyoseridinae (Compositae: Lactuceae). – Bot. J. Linn. Soc. 82: 1–13.

Blanca G. & Díaz de la Guardia C. 1985: Avellara, género nuevo de la tribu Lactuceae (Compositae) en la península Ibérica. – Candollea 40: 447–458.

Bremer K. 1994: Asteraceae. Cladistics and classification. – Portland: Timber.

Kilian N., Gemeinholzer B. & Lack H. W. 2009: Tribe Cichorieae. – Pp. 343–383 In: Funk V. A., Susanna A., Stuessy T. & Bayer R. (ed.), Systematics, evolution, and biogeography of the Compositae. – Vienna: IAPT.

Lack H. W. 2006: Tribe Cichorieae Lam. & DC. – Pp. 180–199 in: Kadereit J. W. & Jeffrey C. (ed.), The families and genera of vascular plants 8. – Berlin: Springer.

Mavrodiev E. V., Edwards C. E., Albach D. C., Gitzendanner M. A., Soltis P. S. & Soltis D. E. 2004: Phylogenetic relationships in subtribe Scorzonerinae (Asteraceae: Cichorioideae: Cichorieae) based on ITS sequence data. – Taxon 53: 699–712.

Stebbins G. L. 1953: A new classification of the tribe Cichorieae, family Compositae. – Madroño 12: 65–81.

Distribution

Africa: Algeria (Algerianative); Canary Is. (Canary Is.native: doubtfully native); Cape Provinces (Eastern Cape Provinceintroduced, Northern Cape Provinceintroduced, Western Cape Provinceintroduced); Egypt (Egyptnative); Ethiopia (Ethiopiaintroduced); Free State (Free Stateintroduced); KwaZulu-Natal (KwaZulu-Natalnative); Lesotho (Lesothointroduced); Libya (Libyanative); Madeira (Madeiranative); Morocconative (Morocconative); Northern Provinces (Gautengintroduced, Mpumalangaintroduced, North-West Provinceintroduced, Northern Provinceintroduced); Tunisia (Tunisianative); Zaire (Zairecultivated) Asia-Temperate: Afghanistannative (Afghanistannative); Altay (Altaynative); Amur (Amurnative); Buryatiya (Buryatiyanative); China North-Central (Beijingintroduced, Gansunative, Hebeinative, Shaanxinative, Shandongnative, Shanxinative); China South-Central (Guizhounative, Hubeinative, Sichuanintroduced, Yunnanintroduced); China Southeast (Anhuinative, Henannative, Jiangsunative); Chita (Chitanative); Cyprus (Cyprusnative); East Aegean Is. (East Aegean Is.native); Gulf States (Bahrainnative, United Arab Emiratesnative); Inner Mongolia (Nei Mongolnative, Ningxianative); Iran (Irannative); Iraq (Iraqnative); Irkutsk (Irkutsknative); Kazakhstan (Kazakhstannative); Khabarovsk (Khabarovsknative); Kirgizistannative (Kirgizistannative); Korea (North Koreanative, South Koreanative); Krasnoyarsk (Krasnoyarsknative); Kuwait (Kuwaitnative); Lebanon-Syria (Lebanonnative, Syrianative); Manchuria (Heilongjiangnative, Jilinnative, Liaoningnative); Mongolianative (Mongolianative); North Caucasusnative (Chechnyanative, Dagestannative, Kabardino-Balkariyanative, Karacheyevo-Cherkessiyanative, Krasnodarnative, Severo-Osetiyanative, Stavropolnative); Oman (Omannative); Palestine (Israelnative, Jordannative); Primorye (Primoryenative); Qinghai (Qinghainative); Sakhalin (Sakhalinnative); Saudi Arabia (Saudi Arabianative); Sinai (Sinainative); Tadzhikistannative (Tadzhikistannative); Tibet (Tibetnative); Transcaucasus (Abkhaziyanative, Adzhariyanative, Armenianative, Azerbaijannative, Georgianative, Nakhichevannative); Turkey (Turkeynative); Turkmenistannative (Turkmenistannative); Tuva (Tuvanative); Uzbekistannative (Uzbekistannative); West Siberia (West Siberianative); Xinjiang (Xinjiangnative); Yakutskiya (Yakutskiyanative); Yemen (North Yemennative, South Yemennative) Asia-Tropical: East Himalaya (Bhutannative); India (Haryananative, Punjabnative, Rajasthannative, Uttar Pradeshnative); Jawa (Jawaintroduced); Myanmar (Myanmarintroduced); Nepal (Nepalnative); Pakistan (Pakistannative); West Himalaya (Himachal Pradeshnative, Jammu-Kashmirnative) Australasia: New South Wales (Australian Capital Territoryintroduced, New South Walesintroduced); New Zealand North (New Zealand Northintroduced); New Zealand South (New Zealand Southintroduced); Northern Territory (Northern Territoryintroduced); Queensland (Queenslandintroduced); South Australia (South Australiaintroduced); Tasmania (Tasmaniaintroduced); Victoria (Victoriaintroduced); Western Australia (Western Australiaintroduced) Europe: Albania (Albanianative); Austrianative (Austrianative, Liechtensteinnative); Baleares (Balearesnative); Baltic States (Estonianative, Kaliningradnative, Latvianative, Lithuanianative); Belarus (Belarusnative); Belgiumnative (Belgiumnative, Luxembourgnative); Bulgaria (Bulgarianative); Central European Russia (Central European Russianative); Corse (Corsenative); Czechoslovakia (Czech Republicnative, Slovakianative); Denmark (Denmarknative); East European Russia (East European Russianative); Finland (Finlandnative); Francenative (Francenative); Germany (Germanynative); Great Britain (Great Britainnative); Greece (Greecenative); Hungary (Hungarynative); Irelandnative (Irelandnative, Northern Irelandnative); Italynative; Kriti (Kritinative); Krym (Krymnative); Netherlands (Netherlandsnative); North European Russia (North European Russianative); Northwest European Russia (Northwest European Russianative); Norway (Norwaynative); Poland (Polandnative); Portugal (Portugalnative); Romania (Romanianative); Sardegna (Sardegnanative); Sicilynative (Maltanative, Sicilynative); South European Russia (South European Russianative); Spainnative (Gibraltarnative, Spainnative); Sweden (Swedennative); Switzerland (Switzerlandnative); Turkey-in-Europe (Turkey-in-Europenative); Ukraine (Moldovanative, Ukrainenative); Yugoslavia (Bosnia-Herzegovinanative, Croatianative, Macedonianative, Montenegronative, Serbianative, Slovenianative) Northern America: Alberta (Albertaintroduced); Arizona (Arizonanative); Arkansas (Arkansasintroduced); British Columbia (British Columbiaintroduced); California (Californiaintroduced); Colorado (Coloradointroduced); Connecticut (Connecticutintroduced); Delaware (Delawareintroduced); District of Columbia (District of Columbiaintroduced); Georgia, U.S.A. (Georgia, U.S.A.native); Idaho (Idahonative); Illinois (Illinoisintroduced); Indiana (Indianaintroduced); Iowa (Iowaintroduced); Kansas (Kansasintroduced); Kentucky (Kentuckyintroduced); Labrador (Labradorintroduced); Louisiana (Louisianaintroduced); Maine (Maineintroduced); Manitoba (Manitobaintroduced); Maryland (Marylandintroduced); Massachusetts (Massachusettsintroduced); Mexico Northwest (Baja Californiaintroduced); Michigan (Michiganintroduced); Minnesota (Minnesotaintroduced); Missouri (Missouriintroduced); Montana (Montanaintroduced); Nebraska (Nebraskaintroduced); Nevada (Nevadaintroduced); New Brunswick (New Brunswickintroduced); New Hampshire (New Hampshireintroduced); New Jersey (New Jerseyintroduced); New Mexico (New Mexicointroduced); New York (New Yorkintroduced); Newfoundland (Newfoundlandintroduced); North Carolina (North Carolinaintroduced); North Dakota (North Dakotaintroduced); Northwest Territories (Northwest Territoriesintroduced); Nova Scotia (Nova Scotiaintroduced); Ohio (Ohiointroduced); Oklahoma (Oklahomaintroduced); Ontario (Ontariointroduced); Oregon (Oregonintroduced); Pennsylvania (Pennsylvaniaintroduced); Prince Edward I. (Prince Edward I.introduced); Québec (Québecintroduced); Rhode I. (Rhode I.introduced); Saskatchewan (Saskatchewanintroduced); South Dakota (South Dakotaintroduced); Tennessee (Tennesseeintroduced); Texas (Texasintroduced); Utah (Utahintroduced); Vermont (Vermontintroduced); Virginia (Virginiaintroduced); Washington (Washingtonnative); West Virginia (West Virginiaintroduced); Wisconsin (Wisconsinintroduced); Wyoming (Wyomingnative); Yukon (Yukonintroduced) Southern America: Argentina Northeast (Buenos Airesintroduced, Córdobaintroduced, La Pampaintroduced); Argentina Northwest (Jujuyintroduced, Mendozaintroduced, San Luisintroduced); Argentina South (Chubutintroduced, Neuquénintroduced, Rio Negrointroduced, Santa Cruzintroduced, Tierra del Fuego (Argentina)introduced); Chile Central (Biobíointroduced, Coquimbointroduced, O'Higginsintroduced, Santiagointroduced, Valparaísointroduced); Chile South (Aisénintroduced); Dominican Republic (Dominican Republicintroduced); Haiti (Haitiintroduced); Puerto Rico (Puerto Ricointroduced)