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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.).
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.
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 (Algeria ‒ native); Canary Is. (Canary Is. ‒ native: doubtfully native); Cape Provinces (Eastern Cape Province ‒ introduced, Northern Cape Province ‒ introduced, Western Cape Province ‒ introduced); Egypt (Egypt ‒ native); Ethiopia (Ethiopia ‒ introduced); Free State (Free State ‒ introduced); KwaZulu-Natal (KwaZulu-Natal ‒ native); Lesotho (Lesotho ‒ introduced); Libya (Libya ‒ native); Madeira (Madeira ‒ native); Morocco ‒ native (Morocco ‒ native); Northern Provinces (Gauteng ‒ introduced, Mpumalanga ‒ introduced, North-West Province ‒ introduced, Northern Province ‒ introduced); Tunisia (Tunisia ‒ native); Zaire (Zaire ‒ cultivated) Asia-Temperate: Afghanistan ‒ native (Afghanistan ‒ native); Altay (Altay ‒ native); Amur (Amur ‒ native); Buryatiya (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 (Chita ‒ native); Cyprus (Cyprus ‒ native); East Aegean Is. (East Aegean Is. ‒ native); Gulf States (Bahrain ‒ native, United Arab Emirates ‒ native); Inner Mongolia (Nei Mongol ‒ native, Ningxia ‒ native); Iran (Iran ‒ native); Iraq (Iraq ‒ native); Irkutsk (Irkutsk ‒ native); Kazakhstan (Kazakhstan ‒ native); Khabarovsk (Khabarovsk ‒ native); Kirgizistan ‒ native (Kirgizistan ‒ native); Korea (North Korea ‒ native, South Korea ‒ native); Krasnoyarsk (Krasnoyarsk ‒ native); Kuwait (Kuwait ‒ native); Lebanon-Syria (Lebanon ‒ native, Syria ‒ native); Manchuria (Heilongjiang ‒ native, Jilin ‒ native, Liaoning ‒ native); Mongolia ‒ native (Mongolia ‒ native); North Caucasus ‒ native (Chechnya ‒ native, Dagestan ‒ native, Kabardino-Balkariya ‒ native, Karacheyevo-Cherkessiya ‒ native, Krasnodar ‒ native, Severo-Osetiya ‒ native, Stavropol ‒ native); Oman (Oman ‒ native); Palestine (Israel ‒ native, Jordan ‒ native); Primorye (Primorye ‒ native); Qinghai (Qinghai ‒ native); Sakhalin (Sakhalin ‒ native); Saudi Arabia (Saudi Arabia ‒ native); Sinai (Sinai ‒ native); Tadzhikistan ‒ native (Tadzhikistan ‒ native); Tibet (Tibet ‒ native); Transcaucasus (Abkhaziya ‒ native, Adzhariya ‒ native, Armenia ‒ native, Azerbaijan ‒ native, Georgia ‒ native, Nakhichevan ‒ native); Turkey (Turkey ‒ native); Turkmenistan ‒ native (Turkmenistan ‒ native); Tuva (Tuva ‒ native); Uzbekistan ‒ native (Uzbekistan ‒ native); West Siberia (West Siberia ‒ native); Xinjiang (Xinjiang ‒ native); Yakutskiya (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 (Jawa ‒ introduced); Myanmar (Myanmar ‒ introduced); Nepal (Nepal ‒ native); Pakistan (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 (New Zealand North ‒ introduced); New Zealand South (New Zealand South ‒ introduced); Northern Territory (Northern Territory ‒ introduced); Queensland (Queensland ‒ introduced); South Australia (South Australia ‒ introduced); Tasmania (Tasmania ‒ introduced); Victoria (Victoria ‒ introduced); Western Australia (Western Australia ‒ introduced) Europe: Albania (Albania ‒ native); Austria ‒ native (Austria ‒ native, Liechtenstein ‒ native); Baleares (Baleares ‒ native); Baltic States (Estonia ‒ native, Kaliningrad ‒ native, Latvia ‒ native, Lithuania ‒ native); Belarus (Belarus ‒ native); Belgium ‒ native (Belgium ‒ native, Luxembourg ‒ native); Bulgaria (Bulgaria ‒ native); Central European Russia (Central European Russia ‒ native); Corse (Corse ‒ native); Czechoslovakia (Czech Republic ‒ native, Slovakia ‒ native); Denmark (Denmark ‒ native); East European Russia (East European Russia ‒ native); Finland (Finland ‒ native); France ‒ native (France ‒ native); Germany (Germany ‒ native); Great Britain (Great Britain ‒ native); Greece (Greece ‒ native); Hungary (Hungary ‒ native); Ireland ‒ native (Ireland ‒ native, Northern Ireland ‒ native); Italy ‒ native; Kriti (Kriti ‒ native); Krym (Krym ‒ native); Netherlands (Netherlands ‒ native); North European Russia (North European Russia ‒ native); Northwest European Russia (Northwest European Russia ‒ native); Norway (Norway ‒ native); Poland (Poland ‒ native); Portugal (Portugal ‒ native); Romania (Romania ‒ native); Sardegna (Sardegna ‒ native); Sicily ‒ native (Malta ‒ native, Sicily ‒ native); South European Russia (South European Russia ‒ native); Spain ‒ native (Gibraltar ‒ native, Spain ‒ native); Sweden (Sweden ‒ native); Switzerland (Switzerland ‒ native); Turkey-in-Europe (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 (Alberta ‒ introduced); Arizona (Arizona ‒ native); Arkansas (Arkansas ‒ introduced); British Columbia (British Columbia ‒ introduced); California (California ‒ introduced); Colorado (Colorado ‒ introduced); Connecticut (Connecticut ‒ introduced); Delaware (Delaware ‒ introduced); District of Columbia (District of Columbia ‒ introduced); Georgia, U.S.A. (Georgia, U.S.A. ‒ native); Idaho (Idaho ‒ native); Illinois (Illinois ‒ introduced); Indiana (Indiana ‒ introduced); Iowa (Iowa ‒ introduced); Kansas (Kansas ‒ introduced); Kentucky (Kentucky ‒ introduced); Labrador (Labrador ‒ introduced); Louisiana (Louisiana ‒ introduced); Maine (Maine ‒ introduced); Manitoba (Manitoba ‒ introduced); Maryland (Maryland ‒ introduced); Massachusetts (Massachusetts ‒ introduced); Mexico Northwest (Baja California ‒ introduced); Michigan (Michigan ‒ introduced); Minnesota (Minnesota ‒ introduced); Missouri (Missouri ‒ introduced); Montana (Montana ‒ introduced); Nebraska (Nebraska ‒ introduced); Nevada (Nevada ‒ introduced); New Brunswick (New Brunswick ‒ introduced); New Hampshire (New Hampshire ‒ introduced); New Jersey (New Jersey ‒ introduced); New Mexico (New Mexico ‒ introduced); New York (New York ‒ introduced); Newfoundland (Newfoundland ‒ introduced); North Carolina (North Carolina ‒ introduced); North Dakota (North Dakota ‒ introduced); Northwest Territories (Northwest Territories ‒ introduced); Nova Scotia (Nova Scotia ‒ introduced); Ohio (Ohio ‒ introduced); Oklahoma (Oklahoma ‒ introduced); Ontario (Ontario ‒ introduced); Oregon (Oregon ‒ introduced); Pennsylvania (Pennsylvania ‒ introduced); Prince Edward I. (Prince Edward I. ‒ introduced); Québec (Québec ‒ introduced); Rhode I. (Rhode I. ‒ introduced); Saskatchewan (Saskatchewan ‒ introduced); South Dakota (South Dakota ‒ introduced); Tennessee (Tennessee ‒ introduced); Texas (Texas ‒ introduced); Utah (Utah ‒ introduced); Vermont (Vermont ‒ introduced); Virginia (Virginia ‒ introduced); Washington (Washington ‒ native); West Virginia (West Virginia ‒ introduced); Wisconsin (Wisconsin ‒ introduced); Wyoming (Wyoming ‒ native); Yukon (Yukon ‒ introduced) Southern America: Argentina Northeast (Buenos Aires ‒ introduced, Córdoba ‒ introduced, La Pampa ‒ introduced); Argentina Northwest (Jujuy ‒ introduced, Mendoza ‒ 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 (Dominican Republic ‒ introduced); Haiti (Haiti ‒ introduced); Puerto Rico (Puerto Rico ‒ introduced)
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