Cichorieae

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Cichorieae

Distribution

Africa: Aldabra native; Algeria nativenative; Angola native; Azores native; Benin native; Botswana introduced; Burkina native; Burundi native; Cameroon native; Canary Is. nativenative; Cape Provinces (Eastern Cape Province native, Northern Cape Province native, Western Cape Province native); Cape Verde nativenative; Central African Republic native; Chad native; Comoros (Comoros native, Mayotte introduced); Congo native; Djibouti native; Egypt native; Equatorial Guinea native; Eritrea native; Ethiopia native; Free State native; Gabon native; Gambia, The introduced; Ghana native; Guinea native; Gulf of Guinea Is. (Bioko native, São Tomé introduced); Ivory Coast native; Kenya native; KwaZulu-Natal native; Lesotho native; Liberia native; Libya native; Madagascar native; Madeira native; Malawi native; Mali native; Mauritania native; Mauritius native; Morocco native (Morocco native); Mozambique native; Namibia native; Niger native; Nigeria native; Northern Provinces (Gauteng native, Mpumalanga native, North-West Province native, Northern Province native); Rodrigues native; Rwanda native; Réunion native; Selvagens native; Senegal native; Seychelles native; Sierra Leone native; Socotra native; Somalia native; St.Helena native; Sudan native; Swaziland native; Tanzania native; Togo native; Tunisia native; Uganda native; Western Sahara nativenative; Zaire native; Zambia native; Zimbabwe native Antarctic: Falkland Is. native Asia-Temperate: Afghanistan nativenative; Altay nativenative; Amur nativenative; Buryatiya native; China North-Central (Beijing native, Gansu native, Hebei native, Shaanxi native, Shandong native, Shanxi native); China South-Central (Chongqing native, Guizhou native, Hubei native, Sichuan native, Yunnan native); China Southeast (Anhui native, Fujian native, Guangdong native, Guangxi native, Henan native, Hong Kong native, Hunan native, Jiangsu native, Jiangxi native, Zhejiang native); Chita native; Cyprus nativenative; East Aegean Is. native; Gulf States (Bahrain native, Qatar native, United Arab Emirates native); Hainan native; Inner Mongolia (Nei Mongol native, Ningxia native); Iran nativenative; Iraq nativenative; Irkutsk native; Japan native (Hokkaido native, Honshu native, Kyushu native, Shikoku native); Kamchatka nativenative; Kazakhstan nativenative; Khabarovsk nativenative; Kirgizistan nativenative; Korea native (North Korea native, South Korea native); Krasnoyarsk native; Kuril Is. nativenative; Kuwait native; Lebanon-Syria native (Lebanon native, Syria native); Magadan nativenative; Manchuria (Heilongjiang native, Jilin native, Liaoning native); Mongolia nativenative; Nansei-shoto native; North Caucasus native (Chechnya native, Dagestan native, Kabardino-Balkariya native, Karacheyevo-Cherkessiya native, Krasnodar native, Severo-Osetiya native, Stavropol native); Ogasawara-shoto native; Oman native; Palestine (Israel native, Jordan native); Primorye nativenative; Qinghai native; Sakhalin nativenative; Saudi Arabia native; Sinai native; Tadzhikistan nativenative; Taiwan native; Tibet nativenative; Transcaucasus (Abkhaziya native, Adzhariya native, Armenia native, Azerbaijan native, Georgia native, Nagorno Karabakh native, Nakhichevan native); Turkey nativenative; Turkmenistan nativenative; Tuva native; Uzbekistan nativenative; West Siberia native; Xinjiang nativenative; Yakutskiya native; Yemen introduced (North Yemen native, South Yemen native) Asia-Tropical: Andaman Is. (Andaman Is. native); Assam (Assam native, Manipur native, Meghalaya native, Mizoram native, Nagaland native); Bangladesh native; Borneo (Sabah native); Cambodia native; East Himalaya native (Arunachal Pradesh native, Bhutan native, Darjiling native, Sikkim native); India introduced (Bihar native, Chattisgarh native, Delhi native, Haryana native, Karnataka native, Kerala native, Madhya Pradesh native, Maharashtra native, Orissa native, Punjab native, Rajasthan native, Tamil Nadu native, Uttar Pradesh native, West Bengal native); Jawa native; Laos nativenative; Malaya (Peninsular Malaysia native); Maldives native; Maluku native; Myanmar nativenative; Nepal native; New Guinea reported in error (Irian Jaya native, Papua New Guinea native); Pakistan nativenative; Philippines native; Sri Lanka native; Sulawesi native; Sumatera native; Thailand native; Vietnam native; West Himalaya native (Himachal Pradesh native, Jammu-Kashmir native, Uttaranchal native) Australasia: Antipodean Is. native; Chatham Is. native; Kermadec Is. introduced; New South Wales (Australian Capital Territory native, New South Wales native); New Zealand North native; New Zealand South native; Northern Territory introduced; Queensland (Queensland native); South Australia native; Tasmania native; Victoria native; Western Australia (Ashmore-Cartier Is. introduced, Western Australia native) Europe: Albania nativenative; Austria native (Austria native, Liechtenstein native); Baleares nativenative; Baltic States native (Estonia native, Kaliningrad native, Latvia native, Lithuania native); Belarus native; Belgium native (Belgium native, Luxembourg native); Bulgaria nativenative; Central European Russia native; Corse nativenative; Czechoslovakia native (Czech Republic native, Slovakia native); Denmark nativenative; East European Russia native; Finland nativenative; France native (Channel Is. native, France native); Føroyar native; Germany nativenative; Great Britain nativenative; Greece nativenative; Hungary native; Iceland native; Ireland native (Ireland native, Northern Ireland native); Italy native (Italy native); Kriti native; Krym native; Netherlands nativenative; North European Russia native; Northwest European Russia native; Norway nativenative; Poland native; Portugal nativenative; Romania nativenative; Sardegna native; Sicily native (Malta native, Sicily native); South European Russia native; Spain native (Andorra native, Gibraltar native, Spain native); Svalbard native; Sweden nativenative; Switzerland nativenative; Turkey-in-Europe native; Ukraine native (Moldova native, Ukraine native); Yugoslavia native (Bosnia-Herzegovina native, Croatia native, Macedonia native, Montenegro native, Serbia native, Slovenia native) Northern America: Alabama native; Alaska native; Alberta native; Aleutian Is. nativenative; Arizona nativenative; Arkansas nativenative; British Columbia native; California nativenative; Colorado native; Connecticut nativenative; Delaware nativenative; District of Columbia nativenative; Florida native; Georgia, U.S.A. nativenative; Greenland native; Idaho native; Illinois nativenative; Indiana nativenative; Iowa nativenative; Kansas native; Kentucky nativenative; Labrador native; Louisiana native; Maine nativenative; Manitoba native; Maryland nativenative; Massachusetts nativenative; Mexican Pacific Is. (Guadalupe I. native); Mexico Central native (Mexico Distrito Federal native, Morelos native, México State native, Puebla native, Tlaxcala native); Mexico Gulf (Veracruz native); Mexico Northeast (Aguascalientes native, Chihuahua native, Coahuila native, Durango native, Guanajuato native, Hidalgo native, Nuevo León native, Querétaro native, San Luis Potosí native, Tamaulipas native, Zacatecas native); Mexico Northwest (Baja California native, Baja California Sur native, Sinaloa native, Sonora native); Mexico Southeast (Campeche introduced, Chiapas native, Quintana Roo introduced, Yucatán introduced); Mexico Southwest (Colima native, Guerrero native, Jalisco native, Michoacán native, Nayarit native, Oaxaca native); Michigan nativenative; Minnesota nativenative; Mississippi native; Missouri nativenative; Montana native; Nebraska native; Nevada native; New Brunswick nativenative; New Hampshire nativenative; New Jersey nativenative; New Mexico nativenative; New York nativenative; Newfoundland (Newfoundland native, St.Pierre-Miquelon native); North Carolina nativenative; North Dakota native; Northwest Territories native; Nova Scotia nativenative; Nunavut native; Ohio nativenative; Oklahoma nativenative; Ontario nativenative; Oregon native; Pennsylvania nativenative; Prince Edward I. nativenative; Québec native; Rhode I. nativenative; Saskatchewan native; South Carolina nativenative; South Dakota native; Tennessee nativenative; Texas native; Utah nativenative; Vermont nativenative; Virginia nativenative; Washington native; West Virginia nativenative; Wisconsin nativenative; Wyoming native; Yukon native Pacific: Cook Is. introduced; Fiji introduced; Hawaii (Hawaiian Is. introduced); New Caledonia introduced South Africa, Republic of: introduced South Sudan: native Southern America: Argentina Northeast (Argentina Distrito Federal native, Buenos Aires native, Chaco native, Corrientes native, Córdoba native, Entre Ríos native, Formosa native, La Pampa native, Misiones native); Argentina Northwest (Catamarca native, Jujuy native, La Rioja native, Mendoza native, Salta native, San Juan native, San Luis native, Santiago del Estero native, Tucuman native); Argentina South (Chubut native, Neuquén native, Rio Negro native, Santa Cruz native, Santa Fé native, Tierra del Fuego (Argentina) native); Aruba introduced; Bahamas native; Belize native; Bermuda native; Bolivia native; Brazil Northeast (Bahia native); Brazil South (Paraná native, Rio Grande do Sul native, Santa Catarina native); Brazil Southeast (Minas Gerais native, Rio de Janeiro native, São Paulo native, Trindade introduced); Cayman Is. introduced; Chile Central native (Biobío native, Coquimbo native, La Araucania native, Maule native, O'Higgins native, Santiago native, Valparaíso native); Chile North native (Antofagasta native, Atacama native, Tarapaca native); Chile South (Aisén native, Los Lagos native, Magellanes native); Colombia native; Costa Rica native; Cuba introducedintroduced; Desventurados Is. native; Dominican Republic native; Ecuador nativenative; El Salvador native; Galápagos introduced; Guatemala nativenative; Haiti (Haiti native, Navassa I. introduced); Honduras native; Jamaica native; Juan Fernández Is. nativenative; Leeward Is. (Anguilla introduced, Antigua-Barbuda introduced, British Virgin Is. native, Guadeloupe introduced, Montserrat introduced, Netherlands Leeward Is. introduced, St.Martin-St.Barthélémy introduced, Virgin Is. introduced); Netherlands Antilles introduced (Bonaire introduced, Curaçao introduced); Nicaragua native; Panamá native; Paraguay native; Peru nativenative; Puerto Rico native; Trinidad-Tobago introduced; Turks-Caicos Is. introduced; Uruguay native; Venezuela native; Venezuelan Antilles introduced; Windward Is. (Barbados introduced, Dominica introduced, Grenada introduced, Martinique introduced, St.Lucia introduced, St.Vincent introduced)

Description

Annual to perennial, acaulescent, scapose or caulescent herbs, more rarely subshrubs, shrubs, rosette shrubs and rosette treelets, sometimes spinescent, exceptionally scandent vines, leaves alternate (except in Shinnersoseris the lower leaves), very rarely spiny (Scolymus, Gundelia), frequently forming a rosette, with latex. Heads solitary or loosely to densely aggregated, sometimes in secondary heads, ligulate (except for Gundelia and Warionia), homogamous, with 1- (exceptionally in Gundelia, in primary heads) or 3-5 to over 600, mostly with a few dozens flowers; receptacle areolate or alveolate, naked, rarely with scales or bristles; involucre cylindric, mostly differentiated into a few imbricate outer series of bracts and a longer inner series, more rarely uniseriate, bracts with or without hyaline margin; corollae with 5-toothed ligule, perfect, predominantly of some shade of yellow, ranging from pale yellow to deep orange-yellow, or of some shade of blue, including whitish so or purple, sometimes also pure white through absence of either the yellowish or bluish colour; anthers basally calcarate and caudate, apical appendage elongate, smooth, filaments smooth; style slender, usually with long, slender branches, sweeping hairs on the shaft and branches; pollen echinolophate or echinate. Achenes cylindrical, or fusiform to slenderly obconoidal, mostly ribbed, sometimes compressed or flattened, apically truncate, attenuate or beaked, often somehow sculptured, mostly glabrous, sometimes papillose or hairy, rarely villose, frequently heteromorphic; pappus of scales or scabrid to barbellate or plumose bristles, sometimes absent.

Common names

Chinese (China): 菊苣族 ju ju zuA; Vietnamese (Vietnam): Tông rau diếpB

Systematics

Circumscription of the Cichorieae

In the traditional circumscription, the Cichorieae were a conveniently recognized tribe, diagnosed by the unique combination of homogamous capitula with 5-dentate, ligulate flowers and the presence of milky latex. Molecular phylogenetic and phylogenomic studies have, however, altered this view. Evidence has been provided that two genera, Gundelia and Warionia, of hitherto uncertain relationships are actually members of the Cichorieae. So far, Gundelia was associated with the Arctotideae (Bremer 1994), Warionia with the Mutisieae or remained unassigned to a tribe (Bremer 1994), or they were treated as the only members of a separate tribe Gundelieae next to Cichorieae (Jeffrey 2006). Both have milky latex but otherwise homogamous capitula with tubular flowers. Hence, by inclusion of these genera the homogamous capitula with 5-dentate, ligulate flowers no longer characterize all the members of the tribe (Kilian & al. 2009).

Neither milky latex nor capitula with only 5-dentate, ligulate flowers represented autapomorphies of the Cichorieae in their traditional sense. Milky latex is otherwise present in some genera of Arctotideae, Cardueae, Liabeae, Mutisieae and Vernonieae (Carlquist 1976), and in a few cases in the Asteroideae (Augier & Mérac 1951; Wagenitz 1976; Bremer 1987, 1994). Milky latex, however, is associated with different anatomical structures in the Compositae. Lactiferous canals, as they are characteristic for all Cichorieae, are otherwise only present in the aerial plant parts of the Arctotideae, whereas other tribes of the Cichorioideae merely have lactiferous cells (Augier & Mérat 1951; Wagenitz 1976). Homogamous capitula with 5-dentate, ligulate flowers are present in a few genera of Mutisieae (Catamixis, Glossarion, Hyaloseris; Bremer 1987, 1994) and, quite evidently by convergent evolution, in the Heliantheae-Coreopsidinae (Fitchia) of subfamily Asteroideae (Carlquist 1957); 5-dentate, ligulate marginal flowers occur in the Vernonieae (Stokesia; Bremer 1987, 1994).

The presence of lactiferous canals in both the subterranean and aerial plant parts seem to be an exclusive feature of the Cichorieae in the present circumscription including Gundelia and Warionia (Augier & Mérac 1951; Wagenitz 1976; Bremer 1987, 1994). Moreover, Gundelia and Warionia share the presence of both (functional) oil ducts and latex canals in the roots (Augier & Mérac 1951) with Scolymus and Scorzonera s.l. (Tieghem 1872; Hoffmann 1890-94; Col 1903-04), while non-functional (relict) oil ducts have been reported from Tragopogon (Tieghem 1885) and Krigia (Holm 1926).

Gundelia, with a much-derived synflorescence of one-flowered capitula aggregated to secondary capitula (syncalathia) of which again a few dozens are aggregated in a second order syncalathium (Classen-Bockhoff & al. 1989), has spiny leaves and pollen similar to Scolymus (Blackmore 1981; Robinson 1994). Gundelia in fact has been shown to form a monophyletic trichotomy with Scolymus and the rest of the Cichorieae by Karis & al. (2001; based on ndhF data). Based on a much broader sampling especially for the basally branching portions of the Cichorieae, Gemeinholzer & al. (in Kilian & al. 2009) and Tremetsberger & al. (2012) have shown that Gundelia forms a clade with Catananche, Hymenonema and Scolymus, which is treated as the subtribe Scolyminae. A phylogenomic study by Mandel & al. (2017) across the Asteraceae confirmed the inclusion of Gundelia in the Cichorieae.

The monospecific Warionia with densely pilose achenes (rare in the Cichorieae but also occurring, e.g., in some species of Scorzonera), has been resolved as sister group to the Cichorieae in their traditional sense in all analyses available so far: Funk & al. (2004), based on trnL-F, ndhF and ITS data, forming a clade together with Gundelia; Goertzen & al. 2003: fig. 3, based on ITS data; Gemeinholzer & al. (in Kilian & al. 2009), based on ITS data; Fu & al. 2016, based on plastid DNA markers. The close relationship to the traditional Cichorieae led Gemeinholzer & Kilian (in Kilian & al. 2009) to establish for Warionia a new subtribe of its own, Warioniinae, within the Cichorieae.

Major clades and recognition of subtribes within the Cichorieae

Whitton & al. (1995) using chloroplast DNA restriction site variation upon 60 Cichorieae taxa were the first to address relationships among major lineages of the tribe and their results agree very well with two more recent attempts by Gemeinholzer & al. (in Kilian & al. 2009), using DNA sequences of the nuclear ITS region and covering 438 taxa, and Tremetsberger & al. (2012), using a subset of 49 species from the former analysis and focusing on divergence time estimations. All analyses revealed similar major lineages.
The analysis by Gemeinholzer & al. (in Kilian & al. 2009) and Tremetsberger & al. (2012) both revealed five major clades, with a total of eleven subclades, within the tribe.
    The first three main clades comprise c. 20 % of the species of the tribe (for species proportions here and in the following Hieracium, Pilosella and Taraxacum not considered):
  • Clade 1 includes solely Warionia (recognized as the subtribe Warioniinae).
  • Clade 2 represents the subtribe Scolyminae.
  • Clade 3 represents the subtribe Scorzonerinae
    The two main clades 4 and 5 comprise roughly 80 % of the species of the tribe:
  • Clade 4 includes five subclades, representing the subtribes Chondrillinae, Crepidinae, Hyoseridinae, Hypochaeridinae and Lactucinae. This main clade comprises 2/3 of the species of the tribe.
  • Clade 5 includes three subclades, representing the subtribes Cichoriinae, Hieraciinae and Microseridinae s.l. This main clade comprises somewhat more than 10 % of the species of the tribe.
The backbone of the tree is essentially unresoved and reconstructed differently by Kilian & al. (2009) and Tremetsberger & al. (2012). In the analysis by Kilian & al. (2009) the main clades 4 and 5 group with the Scolyminae clade, whereas the main clades 4 and 5 group with the Scorzonerinaeclade in that by Tremetsberger & al. (2012). The latter topology with the Scolyminae being sister to the Scorzonerinae + clades 4 and 5 is also corroborated by the analyses of chloroplast restriction sites by Whitton & al.(1994), of chloroplast ndhF sequences by Karis & al. (2001) and by the phylogenomic study by Mandel & al. (2017).

Age of the Cichorieae

The estimated minimum age for the most recent common ancestor of the Cichorieae was revealed as c. 26 (23.2-30.3) mya, indicating that the tribe originated no later than Oligocene, and from the inferred N African-Mediterranean ancestral area of Cichorieae (including Gundelieae, Funk & al. 2005), N Africa seems the most plausible region of origin (Tremetsberger & al. 2012). The clades 4 and 5 diversified no later than Middle/Late Miocene or Early Pliocene (c. 21 (16.8-24.4) mya) (Tremetsberger & al. 2012).
C,D,E,F,G,H,I,J,K,L,M,N,O,P,Q,R,S,T,U,V,W,X,Y

Bibliography

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