Cichoriinae

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Cichoriinae

Distribution

Africa: Algeria native; Angola native; Azores native; Burundi native; Canary Is. native; Cape Provinces (Eastern Cape Province native, Northern Cape Province native, Western Cape Province native); Cape Verde native; Congo cultivated; Egypt native; Eritrea introduced; Ethiopia native; Free State native; Kenya native; KwaZulu-Natal native; Lesotho native; Libya native; Madagascar native; Madeira native; Malawi native; Morocco native; Mozambique native; Northern Provinces (Gauteng native, Mpumalanga native, North-West Province native, Northern Province native); Rwanda native; Socotra native; Somalia native; Sudan native; Swaziland native; Tanzania native; Tunisia native; Uganda native; Zaire native; Zambia native; Zimbabwe native Asia-Temperate: Afghanistan native; Altay introduced; Buryatiya introduced; China North-Central (Beijing introduced, Gansu doubtfully present, Hebei introduced, Shaanxi introduced, Shandong introduced, Shanxi introduced); China Southeast (Henan introduced); Cyprus native; East Aegean Is. native; Gulf States (Qatar native, United Arab Emirates introduced); Iran native; Iraq native; Irkutsk introduced; Kazakhstan native; Khabarovsk introduced; Kirgizistan native; Korea cultivated; Krasnoyarsk introduced; Lebanon-Syria (Lebanon native, Syria native); Manchuria (Heilongjiang introduced, Jilin doubtfully present, Liaoning introduced); Mongolia introduced; North Caucasus native (Chechnya native, Dagestan native, Kabardino-Balkariya native, Karacheyevo-Cherkessiya native, Krasnodar native, Severo-Osetiya native, Stavropol native); Oman introduced; Palestine (Israel native, Jordan native); Primorye introduced; Sakhalin introduced; Saudi Arabia native; Sinai native; Tadzhikistan native; Taiwan introduced; Transcaucasus (Abkhaziya native, Adzhariya native, Armenia native, Azerbaijan native, Georgia native, Nagorno Karabakh native, Nakhichevan native); Turkey native; Turkmenistan native; West Siberia introduced; Xinjiang introduced; Yemen (North Yemen native, South Yemen native) Asia-Tropical: Bangladesh introduced; India (Haryana introduced, Kerala cultivated, Maharashtra introduced, Rajasthan introduced, Uttar Pradesh introduced); Myanmar introduced; Pakistan native; Vietnam cultivated; West Himalaya (Himachal Pradesh introduced) 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 introduced; France native (France introduced); Germany native; Great Britain native; Greece native; Hungary native; Ireland introduced (Ireland introduced, Northern Ireland introduced); Italy native; Kriti native; Krym native; Netherlands native; North European Russia introduced; Northwest European Russia native; Norway introduced; Poland native; Portugal native; Romania native; Sardegna native; Sicily native (Malta native, Sicily native); South European Russia native; Spain 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; Arkansas introduced; British Columbia introduced; California native; Connecticut introduced; Illinois introduced; Indiana introduced; Iowa introduced; Kansas introduced; Labrador introduced; Maine introduced; Manitoba introduced; Massachusetts introduced; Mexico Central (México State introduced); Mexico Northwest (Baja California introduced, Sonora introduced); Michigan introduced; Montana introduced; Nevada introduced; New Brunswick introduced; New Hampshire introduced; New York introduced; Newfoundland (Newfoundland introduced, St.Pierre-Miquelon introduced); North Carolina introduced; Nova Scotia introduced; Ohio introduced; Ontario introduced; Pennsylvania introduced; Prince Edward I. introduced; Québec introduced; Rhode I. introduced; Saskatchewan introduced; Texas introduced; Utah introduced; Vermont introduced Southern America: Argentina Northeast (Argentina Distrito Federal introduced, Buenos Aires introduced, Córdoba introduced, Entre Ríos introduced, La Pampa introduced); Argentina Northwest (Jujuy introduced, Mendoza introduced, Salta introduced, Tucuman introduced); Argentina South (Neuquén introduced, Rio Negro introduced, Santa Fé introduced); Bolivia introduced; Brazil South (Santa Catarina introduced); Chile Central (Biobío introduced, Coquimbo introduced, La Araucania introduced, Maule introduced, O'Higgins introduced, Santiago introduced, Valparaíso introduced); Chile North (Atacama introduced); Chile South (Los Lagos introduced, Magellanes introduced); Cuba introduced; Dominican Republic introduced; El Salvador introduced; Guatemala introduced; Haiti (Haiti introduced); Honduras introduced; Juan Fernández Is. introduced; Puerto Rico introduced; Uruguay introduced; Venezuela introduced; Windward Is. (Martinique introduced)

Systematics

According to the recent molecular phylogenetic analyses by Gemeinholzer & al. (in Kilian & al. 2009) the subtribe Cichoriinae comprises six genera. Four of them, Arnoseris, Cichorium, Rothmaleria and Tolpis, were united already by Stebbins (1953) in his Cichoriinae, together with other, unrelated genera. Later classifications (Jeffrey 1966; Bremer 1994; Lack 2006) split the four genera apart again. To these four genera the analyses by Gemeinholzer & al. added (1) Erythroseris, a genus most recently established for two species from the Horn of Africa and Socotra island formerly placed in Prenanthes (Kilian & Gemeinholzer 2007), and (2) the localized SW North American monospecific Phalacroseris, with oblong-ellipsoid, unbeaked achenes and no pappus, which was revealed to branch off basally to all other taxa of the North American clade (Microseridinae s.l.) by Lee & al. (2003).

The six genera of the Cichoriinae form two subclades. In the first subclade Erythroseris is sister to Cichorium, while Phalacroseris branches off basally to both. In the second subclade Arnoseris is sister to Tolpis, while Rothmaleria branches off basally to both.

Two possible relationships have been suggested in the past for the monospecific SW Mediterranean genus Rothmaleria, namely either with Catananche and Hymenonema (Jeffrey 1966; Bremer 1994; Lack 2006) or with Cichorium (Stebbins 1953; Lack & al. 1980). The latter relationship is not only favoured by the molecular results, it is also supported by similar achenes, the non-aristate paleaceous pappus and the long collecting trichomes of the style (Lack & al. 1980).
The relationship of Arnoseris with Tolpis (Stebbins 1953; Jeffrey 1966; Bremer 1994) rather than with the Hypochaeris alliance (Lack 2006) is morphologically supported by short style branches with short trichomes, corolla tubes covered externally with crisped trichomes and has otherwise been supported by palynological data (Blackmore 1981) featuring small pollen grains with double rows of spines on the equatorial ridges.
The relationship of Tolpis with the Cichorium clade is only revealed in phylogenies of the nuclear marker (nrITS), whereas phylogenies of chloroplast markers reveal Tolpis (and Arnoseris; Gemeinholzer & al., in prep.) to cluster far away from the Cichorium clade (Whitton & al. 1995; Park & al. 2001). The deviating phylogenies of nuclear and the chloroplast markers, with different underlying modes of inheritance, can only be explained by reticulate evolution with an unknown parent preceding generic and species divergence of Tolpis and Arnoseris and resulting in subsequent chloroplast capture. The available molecular analyses have different samplings with respect to ingroup and outgroup taxa and result in slightly different placements of Tolpis, which is, however, most often closely related to Hyoseris and Urospermum. This may indicate former potential hybrid partners. The analyses of the most comprehensive sample was carried out by Gemeinholzer & al. (in prep.), where, in contrast to Park & al. (2001), T. staticifolia as well as T. capensis cluster right within the Tolpis group in both the nuclear and the chloroplast analyses, most likely as result of the broader taxa sampling.


References


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

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

Jeffrey C. 1966: Notes in Compositae I. The Cichorieae in East Tropical Africa. – Kew Bull. 18: 427-486.

Kilian N. & Gemeinholzer B. 2007: Studies in the Compositae of the Arabian Peninsula and Socotra – 7. Erythroseris, a new genus and the previously unknown sister group of Cichorium (Cichorieae subtribe Cichoriinae). – Willdenowia 37: 283-296.

Kilian N., Gemeinholzer B. & Lack H. W. 2009: Tribe Cichorieae. – 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.

Lack H. W., Ern H. & Straka H. 1980: Die Gattung Rothmaleria Font Quer (Asteraceae, Lactuceae). – Willdenowia 10: 37-49.

Lee J., Baldwin B. G. & Gottlieb L. D. 2003: Phylogenetic relationships among the primarily North American genera of Cichorieae (Compositae) based on analysis of 18S-26S nuclear rDNA ITS and ETS sequences. – Syst. Bot. 28: 616-626.

Park S. J., Korompai E. J., Francisco-Ortega J., Santos Guerra A. & Jansen R. K. 2001: Phylogenetic relationships of Tolpis (Asteraceae: Lactuceae) based on ndhF sequence data. – Pl. Syst. Evol. 226: 23-33.

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

Whitton J., Wallace R.S. & Jansen R.K. 1995: Phylogenetic relationships and patterns of character change in the tribe Lactuceae (Asteraceae) based on chloroplast DNA restriction site variation. – Canad. J. Bot. 73: 1058-1073.