Crepidinae

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Crepidinae

Systematics

The Crepidinae as redefined by Gemeinholzer & al. (in Kilian & al. 2009) comprise c. 360 species (the Taraxacum microspecies not counted) or more than 1/4 of the species of the tribe, making it the largest subtribe of the Cichorieae. The Chondrilla alliance has been removed from the Crepidinae as a separate subtribe Chondrillinae (see there).

A comprehensive molecular phylogeny of the subtribe is still not available, but the initial analyses in Kilian & al. (2009) and by Wang & al. (2020), together with a number of studies focussing on single genera or generic alliances allow to identify a number of fairly well supported clades, while the relationships between these clades remain more uncertain so far.

In the Crepis-Lapsana-Rhagadiolus clade, Crepis species of Babcock’s sections Intybellia, Lagoseris, Microcephalum, Phaecasium and Pterotheca cluster in the nuclear and plastid analyses by Enke & Gemeinholzer (2008) in a clade that is the sister group to Lapsana and Rhagadiolus, two genera formerly placed in the Hypochaeridinae (Bremer 1994, Lack 2006, but with affinities to the Crepidinae already shown by Whitton & al 1995). This part of Crepis includes, but is larger than, the Crepis segregate Lagoseris of some authors. The major part of Crepis forms the sister group to the former subclade. The monospecific afroalpine Dianthoseris was found by Enke & al. (2008) to be nested right into, and to form a congener of, this Crepis s.str. Morphologically, however, both parts of Crepis are ill-delimited from each other, which speaks strongly against recognition of Lagoseris as a separate genus besides Crepis (Enke & Gemeinholzer 2008). Since Lapsana and Rhagadiolus, in contrast, are morphologically well delimited from Crepis s.l., it appears, at least for the time being, the best solution to maintain Lapsana and Rhagadiolus and accept Crepis as a paraphyletic genus.

The Crepidiastrum-Lapsanastrum-Youngia clade (Deng & al. 2014, Kilian & al. 2009, see also Liu & al. 2013) is perhaps sister to the Crepis-Lapsana-Rhagadiolus clade. Crepidiastrum is included in the wider sense as was first established by Shih & Kilian (in Shih & al. 2011) and largely confirmed by Peng & al. (2014), thus embracing the former genera Paraixeris and Crepidifolium (Sennikov & Illarionova 2008). To this Youngia forms the sister group (Zhang & al. 2011, Peng & al. 2014, Deng & al. 2014), while Lapsanastrum (Pak & Bremer 1995), as revealed by Deng & al. (2014) is deeply nested within Youngia, consituting some sort of a parallel case to Lapsana and Rhagadiolus being nested in Crepis.

The Ixeris-Ixeridium-Taraxacum clade (Enke & Gemeinholzer 2008, Zhang & al. 2011, Nakamura & al. 2014) includes the genera Ixeris and Ixeridium in the circumscription by Pak & Kawano (1992), which was confirmed by Shih & Kilian (in Shih & al. 2011) and Nakamura & al. 2014. They have a sister group relationship. In turn Taraxacum appeares to be sister to this clade. Moreoever, this clade may perhaps also include Askellia according to the analysis by Zhang & al. (2011) and Liu & al. (2013). In the analyses by Enke & Gemeinholzer (2008) and Peng & al. (2014), however, Askellia appears as sister to Crepis s.l. Askellia, the former Crepis sect. Ixeridopsis, which has been recognised as a separate genus also by Sennikov & Illarionova (2008) on morphological ground, was first shown in the molecular phylogenetic analysis by Enke & Gemeinholzer (2008) to constitute a genus separate from Crepis. Already Babcock (1947) stated the morphologically intermediate position of Askellia between Crepis and Ixeris, but most of the species (e.g., A. nana, A. flexuosa), used to be treated under Youngia, to which they were reassigned by Adylov & Zuckerwanik (1993). However, Askellia has terete achenes and a basic chromosome number of x = 7 (otherwise not present in Crepis), while the achenes of Youngia are compressed and angular and the chromosome number is x = 8. Also no molecular analysis revealed an affinity with Youngia.

The Garhadiolus-Heteracia clade (Kilian & al. 2009; Zhang & al. 2011; Kilian & al. 2017; Wang & al. 2020) includes four small or monotypic genera, Garhadiolus, Heteracia, Heteroderis and Lagoseriopsis and seems to be sister to the large and diverse Dubyaea-Nabalus-Soroseris-Syncalathium clade. Confirmed (see Whitton & al. 1995) is thus the inclusion of Garhadiolus, formerly placed in the Hypochaeridinae (Bremer 1994; Lack 2006), in the Crepidinae.

The large Dubyaea-Nabalus-Soroseris-Syncalathium clade (Kilian & al. 2009, Zhang & al. 2011; Liu & al. 2013; Wang & al. 2020) is predominantly Asian (extending into North America) and still lacks resolution. Nabalus and Syncalathium were formerly placed in the Lactucinae (Bremer 1994; Lack 2006). This clade, moreover, includes Hololeion, formerly placed in the Hieraciinae (Bremer 1994; Lack 2006). Syncalathium is actually diphylectic; in the sense of the type of the name, which is provided by S. sukaczevii (correctly to be named S. kawaguchii), the genus is a member of the Crepidinae, whereas S. souliei is nested in the Lactuca alliance, a result also corroborated by karyological data (Zhang & al. 2007, 2009). The recognition of Nabalus as a genus separate from Prenanthes, including all North American and several Central and East Asian members of the latter genus, confirms Stebbins (1940: 63). He had concluded from studies of the achene vascularisation that the species of Nabalus are much closer to Dubyaea and Soroseris than to Prenanthes purpurea, which provides the type of the name Prenanthes. Nabalus is, however, polyphyletic: the Chinese Nabalus species group in a different subclade than the North American N. altissimus and N. trifoliolatus, the latter providing the type of the generic name Nabalus, and the northwestern North American N. sagittatus is sister to the former two clades (Zhang & al. 2011; Kilian & al. 2017). A further member is Stebbinsia (Zhang & al. 2011) with its only species S. umbrella but this is nested right within Soroseris, as was already assumed by Lack (2007) from morphology. Dubyaea in its current circumscription also appears to be polyphyletic (Liu & al. 2013). Sonchella, accommodating two Central to E Asian species, and Sinoseris, ecologically outstanding for its distribution in the subtropical (warm-temperate) highlands of Sichuan and Yunnan, are also a member of the Dubyaea-Nabalus-Soroseris-Syncalathium clade (Kilian & al. 2017; Wang & al. 2020).
A,B,C,D,E,F,G,H,I,J,K,L,M,N,O,P,Q,R,S,T,U,V

Distribution

Africa: Algeria native; Angola native; Azores native; Burundi native; Cameroon native; Canary Is. nativenative; Cape Provinces (Eastern Cape Province native, Northern Cape Province introduced, Western Cape Province introduced); Comoros (Comoros introduced); Djibouti introduced; Egypt native; Eritrea native; Ethiopia native; Free State native; Kenya native; KwaZulu-Natal native; Lesotho native; Libya native; Madagascar introduced; Madeira native; Malawi native; Mauritius introduced; Morocco native (Morocco native); Mozambique native; Namibia introduced; Nigeria native; Northern Provinces (Gauteng native, Mpumalanga native, North-West Province native, Northern Province native); Rodrigues introduced; Rwanda native; Réunion introduced; Selvagens native; Seychelles introduced; Somalia native; Sudan native; Swaziland native; Tanzania native; Tunisia native; Uganda native; 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, Hunan native, Jiangsu native, Jiangxi native, Zhejiang native); Chita native; Cyprus nativenative; East Aegean Is. native; Gulf States (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 native; Khabarovsk nativenative; Kirgizistan native; Korea native (North Korea native, South Korea native); Krasnoyarsk native; Kuril Is. nativenative; 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 native; Transcaucasus (Abkhaziya native, Adzhariya native, Armenia native, Azerbaijan native, Georgia 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: Assam (Assam native, Manipur native, Meghalaya native, Nagaland native); Bangladesh native; Borneo (Sabah native); Cambodia native; East Himalaya native (Arunachal Pradesh native, Bhutan native, Darjiling native, Sikkim native); India (Bihar native, Delhi native, Haryana native, Kerala native, Maharashtra native, Punjab native, Tamil Nadu native, Uttar Pradesh native, West Bengal native); Jawa native; Laos native; Malaya (Peninsular Malaysia native); Maluku native; Myanmar nativenative; Nepal native; New Guinea (Irian Jaya native, Papua New Guinea native); Pakistan nativenative; Philippines native; Sri Lanka native; Sulawesi 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 (Western Australia native) 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 native; France native (Channel Is. native, France native); Føroyar native; Germany native; Great Britain nativenative; Greece native; Hungary native; Iceland native; Ireland native (Ireland native, Northern Ireland native); Italy native (Italy native); Kriti native; Krym native; Netherlands native; North European Russia native; Northwest European Russia native; Norway native; Poland native; Portugal native; Romania native; Sardegna native; Sicily native (Malta native, Sicily native); South European Russia native; Spain native (Andorra native, Gibraltar native, Spain native); Svalbard 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: Alabama native; Alaska native; Alberta native; Aleutian Is. nativenative; Arizona native; Arkansas native; British Columbia native; California native; Colorado native; Connecticut native; Delaware native; District of Columbia native; Florida native; Georgia, U.S.A. native; Greenland native; Idaho native; Illinois native; Indiana native; Iowa native; Kansas native; Kentucky native; Labrador native; Louisiana native; Maine native; Manitoba native; Maryland native; Massachusetts native; Mexico Central (Mexico Distrito Federal native, México State native, Puebla native); Mexico Gulf (Veracruz native); Mexico Northeast (Chihuahua native, Coahuila native, Durango native, Hidalgo native, San Luis Potosí native, Tamaulipas native); Mexico Northwest (Sonora native); Mexico Southeast (Chiapas native); Mexico Southwest (Michoacán introduced, Oaxaca native); Michigan native; Minnesota native; Mississippi native; Missouri native; Montana native; Nebraska native; Nevada native; New Brunswick native; New Hampshire native; New Jersey native; New Mexico native; New York native; Newfoundland (Newfoundland native, St.Pierre-Miquelon native); North Carolina native; North Dakota native; Northwest Territories native; Nova Scotia native; Nunavut native; Ohio native; Oklahoma native; Ontario native; Oregon native; Pennsylvania native; Prince Edward I. native; Québec native; Rhode I. native; Saskatchewan native; South Carolina native; South Dakota native; Tennessee native; Texas native; Utah native; Vermont native; Virginia native; Washington native; West Virginia native; Wisconsin native; 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 introduced, Buenos Aires introduced, Corrientes introduced, Córdoba introduced, Entre Ríos native, Formosa introduced, La Pampa introduced, Misiones introduced); Argentina Northwest (Catamarca introduced, Jujuy introduced, La Rioja native, Mendoza native, Salta native, San Juan native, San Luis introduced, Santiago del Estero introduced, Tucuman introduced); Argentina South (Chubut native, Neuquén native, Rio Negro native, Santa Cruz native, Santa Fé introduced, Tierra del Fuego (Argentina) native); Bahamas introduced; Bermuda native; Bolivia introduced; Brazil South (Paraná native, Rio Grande do Sul introduced, Santa Catarina native); Cayman Is. introduced; Chile Central native (Biobío native, Coquimbo introduced, La Araucania introduced, Maule introduced, O'Higgins introduced, Santiago native, Valparaíso introduced); Chile North (Antofagasta introduced, Atacama introduced); Chile South (Aisén introduced, Los Lagos native, Magellanes native); Colombia native; Costa Rica native; Cuba introducedintroduced; Dominican Republic native; Ecuador introduced; El Salvador introduced; Guatemala native; Haiti (Haiti native, Navassa I. introduced); Honduras native; Jamaica native; Juan Fernández Is. native; Leeward Is. (Virgin Is. introduced); Netherlands Antilles introduced; Nicaragua introduced; Panamá introduced; Paraguay introduced; Peru native; Puerto Rico introduced; Uruguay introduced; Venezuela native; Windward Is. (Barbados introduced, Dominica introduced)

Credits

Zhang Jianwen 2009: Systematics (data).

Bibliography

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