Tragopogon

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Common names

Albanian (Albania): LulebrigjeA,1; Arabic (Lebanon): سَلْسَفيلB,1, فوميB,1; Arabic (Syria): سَلْسَفيلC,1, فوميC,1; Armenian (Armenia): ԱյծեմորուսD,1, ՍինձD,1; Bulgarian (Bulgaria): Козя брадаE,1; Chinese (China): 婆罗门参属 po luo men shen shuF; Estonian (Estonia): PiimjuurG,1; French (Corse): SalsifisH,1; German (Germany): BocksbartI,1; Hebrew (Israel): זְקַן-תַּיִשׁJ,1, זקן-תישJ,1; Italian (Italy): Barba di BeccoK,1; Latvian (Latvia): PlostbārdisG,1; Lithuanian (Lithuania): PūtelisG,1; Polish (Poland): KozibródL,1; Romanian (Moldova): Барба кэприйM,1; Russian (Armenia): КозлобородникD,1; Swedish (Sweden): HaverrotN,1; Turkish (Turkey): YemlıkO
1. recommended

Description

Herbs, perennial, biennial, or rarely annual, glabrous or tomentulose to floccose [or lanate] especially at leaf bases and below capitula, often glabrescent; if biennial then vertical roots spindle-shaped; if perennial then with well-developed caudex. Stem simple or sparingly branched. Leaves basal and cauline, sessile, linear, lanceolate, or narrowly oblong, margin entire or undulate. Capitulum terminal, solitary or sometimes capitula few to many, large, with (20–)40–180 or more florets; peduncle often apically inflated and normally without bracts. Involucre cylindric at anthesis, mostly 10–20 mm or more in diam. Phyllaries 5–15(or 16), in 1(or 2) row(s), linear-lanceolate, triangular-lanceolate, or linear, ± equal, abaxially glabrous [with intertwining hairs], margins white and narrowly pellucid, apex acute. Receptacle naked. Florets with ligules yellow, mauve, orange, purple, or violet. Achene dark to pale brown, straw-colored, or whitish, heteromorphic, outer ones ± cylindric or curviform (fusiform), central ones cylindric and less tuberculate or smooth; achene body normally tuberculate, with 5 fairly well-differentiated ribs, with or rarely without hollows in pericarp; beak slender or stout if present; pappus disk ± pubescent. Pappus persistent, dirty white, yellowish, or slightly fulvous; awns 12–20 or more in 1 or 2 rows, unequal, softly fimbriately plumose, apically scabrid.

from: Shih C., Sukhorukov A. P. & Mavrodiev E. V. in Wu Z. Y. & al. (ed.), Flora of China 20–21: 207–208. 2011, Beijing & St Louis.

Systematics

Tragopogon is an Old World genus of c. 150 species, which is distributed across Europe and temperate Asia from the Atlantic to the Pacific Ocean as well as in N Africa, having a centre of distribution in E Europe and W Asia (Mavrodiev & al. 2005; Bell & al. 2012). Delimitation between species is sometimes difficult and molecular studies also indicated the existence of cryptic species, in particular among the widespread T. porrifolius, T. dubius and T. pratensis, hence the number of species in the genus is perhaps even higher (Mavrodiev & al. 2007, 2008). The genus has been introduced to North America in historical times and occurs there with about 10 species; two of them, T. mirus and T. miscellus, are native and restricted to North America, and are of allopolyploid origin from introduced European parental species (Ownbey 1950; Soltis & al. 2004; Malinska & al. 2011).

Tragopogon includes biennial and perennial herbs, which are easily recognised by their grass-like, linear or linear-lanceolate leaves, simple, or sparingly branched stems with correspondingly only one or a few capitula only, involucres with a single row of phyllaries, receptacles without scales, achenes with a softly fimbriately plumose pappus and a basic chromosome number of x = 6.

The monophyly of the genus is strongly supported by several phylogenetic analyses based on nuclear markers (Marvodiev & al. 2004; 2005, 2012; Bell & al. 2012). The monospecifc genus Geropogon is demonstrated by Mavrodiev & al. (2012) to be the immediate sister to Tragopogon, whereas that genus appeared with low support as sister to Podospermum in a previous analysis (Mavrodiev & al. 2004). Geropogon is morphologically very similar to and was sometimes included in Tragopogon, but differs in particular by a dimorphic pappus (plumose in inner achenes of a capitulum, of 5 simply hairs in outer achenes) and a basic chromosome number of x = 7 (Mavrodiev & al. 2004, 2012).

The history of the systematics of Tragopogon is briefly outlined by Mavrodiev & al. (2005). The hitherto most extensive system of classification was provided by Borisova (1964) in the frame of Flora USSR, including 79 species. Rechinger (1977) extended her treatment to Flora Iranica and other authors partly modified Borisova's treatment (see Mavrodiev & al. 2005). Molecular phylogenetic analyses (Mavrodiev & al. 2005, 2012) revealed some clades corresponding to sections recognised by Borisova (1964), thus confirming them as largely monophyletic (T. sect. Tragopogon, Brevirostris, Hebcarpus and Chromopappus), while other sections (T. sect. Angustissimi, Majores, Collini and Profundisulcati) are apparently not monophyletic.

Relationships among the clades recognised in Tragopogon as well as among species within these clades remain largely unresolved, in spite of the use of seven nuclear loci with over 6900 bp in the most recent analysis by Mavrodiev & al. (2012). A possible explanation is a rapid diversification following the origin of Tragopogon. A simultaneous estimate of phylogeny and divergence times of Tragopogon by Bell & al. (2012) in fact demonstrates the age of a major split and subsequent rapid divergence within Tragopogon to be c. 2.6 Ma (1.7–5.4 Ma using various clock estimates) and a diversification rate of about 0.84–2.71 species/Myr for the crown group. The latter is comparable to that for the rapid Eurasian radiation in Dianthus, which often co-occurs with Tragopogon in the latter's centre of diversification (Bell & al. 2012). The inferred diversification time follows the Messinian Salinity Crisis in the late Miocene, between 5.3 and 5.96 Ma, during which a temporary isolation of the Mediterranean Sea from the Atlantic Ocean resulted in the gradual drying of the Mediterranean Basin and a climate shift in the region from a warm, humid climate to a seasonality with summer droughts and cold, humid winters, under which dry meadow communities developed (Bell & al. 2012).

References


Bell C. D., Mavrodiev E. V., Soltis P. S., Calaminus A. K., Albach D. C., Cellinese N., Garcia-Jacas N. & Soltis D. E. 2012: Rapid diversification of Tragopogon and ecological associates in Eurasia. – J. Evol. Biol. 25: 2470–2480.

Borisova A. G. 1964: Tragopogon. – Pp. 115–196 in: Bobrov E. G. & Tzvelev N. N. (ed.), Flora URSS 29. – Mosqua & Leningrad: Nauka.

Malinska H., Tate J. A., Mavrodiev E., Matyasek R., Lim K. Y., Leitch A. R., Soltis D. E., Soltis P. S. & Kovarik A. 2011: Ribosomal RNA genes evolution in Tragopogon: a story of New and Old World allotetraploids and the synthetic lines. – Taxon 60: 348–354.

Mavrodiev E. V., Edwards C. E., Albach D. E., 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.

Mavrodiev E. V., Tancig M., Sherwood A. M., Gitzendanner M. A., Rocca J., Soltis P. S., Soltis D. E. 2005: Phylogeny of Tragopogon L. (Asteraceae) based on internal and external transcribed spacer sequence data. – Int. J. Pl. Sci. 166: 117–133.

Mavrodiev E. V., Soltis P. S., Glitzendanner M. A., Baldini R. M. & Soltis D. E. 2007: Polyphyly of Tragopogon porrifolius L. (Asteraceae), a European native with intercontinental disjuncts. – Int. J. Pl. Sci. 168: 889–904.

Mavrodiev E. V., Soltis P. S. & Soltis D. E. 2008: Putative parentage of six Old World polyploids in Tragopogon L. (Asteraceae: Scorzonerinae) based on ITS, ETS, and plastid sequence data. – Taxon 57: 1215–1232.

Mavrodiev E. V., Gitzendanner M., Calaminus A. K., Baldini R. M., Soltis P. S. & Soltis D. E. 2012: Molecular phylogeny of Tragopogon L. (Asteraceae) based on seven nuclear loci (Adh, GapC, LFY, AP3, PI, ITS, and ETS). – Webbia 67: 111–137.

Ownbey M. 1950: Natural hybridization and amphiploidy in the genus Tragopogon. – Amer. J. Bot. 37: 487–499.

Rechinger K. H. 1977: Tragopogon. – Pp. 83–120 in: Rechinger K. H. (ed.), Flora Iranica 122. – Graz: Akademische Druck und Verlagsanstalt.

Soltis D. E., Soltis P. S., Pires J. C., Kovarik A. & Tate J. 2004: Recent and recurrent polyploidy in Tragopogon (Asteraceae): genetics, genomic, and cytogenetic comparisons. – Biol. J. Linn. Soc. 82: 485–501.

Distribution

Africa: Algeria (Algerianative); Canary Is. (Canary Is.native: doubtfully native); Cape Provinces (Eastern Cape Provinceintroduced, Northern Cape Provinceintroduced, Western Cape Provinceintroduced); Ethiopia (Ethiopiaintroduced); Free State (Free Stateintroduced); KwaZulu-Natal (KwaZulu-Natalintroduced); Lesotho (Lesothointroduced); Libya (Libyanative); Morocconative (Morocconative); Northern Provinces (Gautengintroduced, Mpumalangaintroduced, North-West Provinceintroduced, Northern Provinceintroduced); Tunisia (Tunisianative) Asia-Temperate: Afghanistan (Afghanistannative); Altay (Altaynative); Amur (Amurintroduced); Buryatiya (Buryatiyanative); China North-Central (Beijingintroduced, Shaanxiintroduced); China South-Central (Guizhouintroduced, Sichuanintroduced, Yunnanintroduced); Chita (Chitanative); Cyprus (Cyprusnative); East Aegean Is. (East Aegean Is.native); Iran (Irannative); Iraq (Iraqnative); Irkutsk (Irkutsknative); Kazakhstan (Kazakhstannative); Khabarovsk (Khabarovskintroduced); Kirgizistan (Kirgizistannative); Krasnoyarsk (Krasnoyarsknative); Lebanon-Syria (Lebanonnative, Syrianative); Mongolianative (Mongolianative); North Caucasusnative (Chechnyanative, Dagestannative, Kabardino-Balkariyanative, Karacheyevo-Cherkessiyanative, Krasnodarnative, Severo-Osetiyanative, Stavropolnative); Palestine (Israelnative, Jordannative); Primorye (Primoryeintroduced); Qinghai (Qinghainative); Saudi Arabia (Saudi Arabiaintroduced); Sinai (Sinainative); Tadzhikistan (Tadzhikistannative); Tibet (Tibetnative); Transcaucasus (Abkhaziyanative, Adzhariyanative, Armenianative, Azerbaijannative, Georgianative, Nakhichevannative); Turkey (Turkeynative); Turkmenistan (Turkmenistannative); Tuva (Tuvanative); Uzbekistan (Uzbekistannative); West Siberia (West Siberianative); Xinjiang (Xinjiangnative) Asia-Tropical: East Himalaya (Bhutannative); India (Uttar Pradeshnative); 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 Russiaintroduced); 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)

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