Crepidinae

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Distribution

Africa: Algeria (Algerianative); Angola (Angolanative); Azores (Açôresnative); Burundi (Burundinative); Cameroon (Cameroonnative); Canary Is. (Canary Is.native); Cape Provinces (Eastern Cape Provincenative, Western Cape Provinceintroduced); Comoros (Comorosintroduced); Djibouti (Djiboutiintroduced); Egypt (Egyptnative); Eritrea (Eritreanative); Ethiopia (Ethiopianative); Free State (Free Statenative); Kenya (Kenyanative); KwaZulu-Natal (KwaZulu-Natalnative); Lesotho (Lesothonative); Libya (Libyanative); Madagascar (Madagascarintroduced); Madeira (Madeiranative); Malawi (Malawinative); Mauritius (Mauritiusintroduced); Morocconative; Mozambique (Mozambiquenative); Namibia (Namibiaintroduced); Nigeria (Nigerianative); Northern Provinces (Gautengnative, Mpumalanganative, North-West Provincenative, Northern Provincenative); Rodrigues (Rodriguesintroduced); Rwanda (Rwandanative); Réunion (Réunionintroduced); Selvagens (Selvagensnative); Seychelles (Seychellesintroduced); Somalia (Somalianative); Sudan (Sudannative); Swaziland (Swazilandnative); Tanzania (Tanzanianative); Tunisia (Tunisianative); Uganda (Ugandanative); Zaire (Zairenative); Zambia (Zambianative); Zimbabwe (Zimbabwenative) Asia-Temperate: Afghanistannative (Afghanistannative); Altay (Altaynative); Amurnative (Amurnative); Buryatiya (Buryatiyanative); China North-Central (Beijingnative, Gansunative, Hebeinative, Shaanxinative, Shandongnative, Shanxinative); China South-Central (Chongqingnative, Guizhounative, Hubeinative, Sichuannative, Yunnannative); China Southeast (Anhuinative, Fujiannative, Guangdongnative, Guangxinative, Henannative, Hunannative, Jiangsunative, Jiangxinative, Zhejiangnative); Chita (Chitanative); Cyprusnative (Cyprusnative); East Aegean Is. (East Aegean Is.native); Gulf States (United Arab Emiratesnative); Hainan (Hainannative); Inner Mongolia (Nei Mongolnative, Ningxianative); Irannative (Irannative); Iraqnative (Iraqnative); Irkutsk (Irkutsknative); Japannative (Hokkaidonative, Honshunative, Kyushunative, Shikokunative); Kamchatkanative (Kamchatkanative); Kazakhstan (Kazakhstannative); Khabarovsknative (Khabarovsknative); Kirgizistan (Kirgizistannative); Koreanative (North Koreanative, South Koreanative); Krasnoyarsk (Krasnoyarsknative); Kuril Is.native (Kuril Is.native); Lebanon-Syria (Lebanonnative, Syrianative); Magadannative (Magadannative); Manchuria (Heilongjiangnative, Jilinnative, Liaoningnative); Mongolianative (Mongolianative); Nansei-shoto (Nansei-shotonative); North Caucasusnative (Chechnyanative, Dagestannative, Kabardino-Balkariyanative, Karacheyevo-Cherkessiyanative, Krasnodarnative, Severo-Osetiyanative, Stavropolnative); Ogasawara-shoto (Ogasawara-shotonative); Oman (Omannative); Palestine (Israelnative, Jordannative); Primoryenative (Primoryenative); Qinghai (Qinghainative); Sakhalinnative (Sakhalinnative); Saudi Arabia (Saudi Arabianative); Sinai (Sinainative); Tadzhikistan (Tadzhikistannative); Taiwan (Taiwannative); Tibet (Tibetnative); Transcaucasus (Abkhaziyanative, Adzhariyanative, Armenianative, Azerbaijannative, Georgianative, Nakhichevannative); Turkey (Turkeynative); Turkmenistannative (Turkmenistannative); Tuva (Tuvanative); Uzbekistan (Uzbekistannative); West Siberia (West Siberianative); Xinjiang (Xinjiangnative); Yakutskiya (Yakutskiyanative); Yemenintroduced (North Yemennative, South Yemennative) Asia-Tropical: Assam (Assamnative, Manipurnative, Meghalayanative, Nagalandnative); Bangladesh (Bangladeshnative); Borneo (Sabahnative); Cambodia (Cambodianative); East Himalaya (Arunachal Pradeshnative, Bhutannative, Darjilingnative, Sikkimnative); India (Biharnative, Delhinative, Haryananative, Keralanative, Orissanative, Punjabnative, Tamil Nadunative, Uttar Pradeshnative, West Bengalnative); Jawa (Jawanative); Laos (Laosnative); Malaya (Peninsular Malaysianative); Maluku (Malukunative); Myanmarnative (Myanmarnative); Nepal (Nepalnative); New Guinea (Irian Jayanative, Papua New Guineanative); Pakistannative (Pakistannative); Philippines (Philippinesnative); Sri Lanka (Sri Lankanative); Sulawesi (Sulawesinative); Thailand (Thailandnative); Vietnamnative (Vietnamnative); West Himalayanative (Himachal Pradeshnative, Jammu-Kashmirnative, Uttaranchalnative) Australasia: Chatham Is. (Chatham Is.introduced); Kermadec Is. (Kermadec Is.introduced); New South Wales (Australian Capital Territoryintroduced, New South Walesnative); New Zealand North (New Zealand Northintroduced); New Zealand South (New Zealand Southintroduced); Queensland (Queenslandnative); 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 (Channel Is.native, Francenative); Føroyar (Føroyarnative); Germany (Germanynative); Great Britain (Great Britainnative); Greece (Greecenative); Hungary (Hungarynative); Iceland (Icelandnative); Irelandnative (Irelandnative, Northern Irelandnative); Italynative (Italynative); Kriti (Kritinative); Krym (Krymnative); Netherlands (Netherlandsnative); North European Russia (North European Russianative); 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 (Andorranative, Gibraltarnative, Spainnative); Svalbard (Svalbardnative); Sweden (Swedennative); Switzerland (Switzerlandnative); Turkey-in-Europe (Turkey-in-Europenative); Ukraine (Moldovanative, Ukrainenative); Yugoslavia (Bosnia-Herzegovinanative, Croatianative, Macedonianative, Montenegronative, Serbianative, Slovenianative) Northern America: Alabama (Alabamanative); Alaska (Alaskanative); Alberta (Albertanative); Arizona (Arizonanative); Arkansas (Arkansasnative); British Columbia (British Columbianative); California (Californianative); Colorado (Coloradonative); Connecticut (Connecticutnative); Delaware (Delawarenative); District of Columbia (District of Columbianative); Florida (Floridanative); Georgia, U.S.A. (Georgia, U.S.A.native); Greenland (Greenlandnative); Idaho (Idahonative); Illinois (Illinoisnative); Indiana (Indiananative); Iowa (Iowanative); Kansas (Kansasnative); Kentucky (Kentuckynative); Labrador (Labradornative); Louisiana (Louisiananative); Maine (Mainenative); Manitoba (Manitobanative); Maryland (Marylandnative); Massachusetts (Massachusettsnative); Mexico Central (México Stateintroduced); Mexico Northeast (Chihuahuanative); Michigan (Michigannative); Minnesota (Minnesotanative); Mississippi (Mississippinative); Missouri (Missourinative); Montana (Montananative); Nebraska (Nebraskanative); Nevada (Nevadanative); New Brunswick (New Brunswicknative); New Hampshire (New Hampshirenative); New Jersey (New Jerseynative); New Mexico (New Mexiconative); New York (New Yorknative); Newfoundland (Newfoundlandnative, St.Pierre-Miquelonnative); North Carolina (North Carolinanative); North Dakota (North Dakotanative); Northwest Territories (Northwest Territoriesnative); Nova Scotia (Nova Scotianative); Nunavut (Nunavutnative); Ohio (Ohionative); Oklahoma (Oklahomanative); Ontario (Ontarionative); Oregon (Oregonnative); Pennsylvania (Pennsylvanianative); Prince Edward I. (Prince Edward I.native); Québec (Québecnative); Rhode I. (Rhode I.native); Saskatchewan (Saskatchewannative); South Carolina (South Carolinanative); South Dakota (South Dakotanative); Tennessee (Tennesseenative); Texas (Texasnative); Utah (Utahnative); Vermont (Vermontnative); Virginia (Virginianative); Washington (Washingtonnative); West Virginia (West Virginianative); Wisconsin (Wisconsinnative); Wyoming (Wyomingnative); Yukon (Yukonnative) Pacific: Fiji (Fijiintroduced); Hawaii (Hawaiian Is.introduced); New Caledonia (New Caledoniaintroduced) Southern America: Argentina Northeast (Argentina Distrito Federalintroduced, Buenos Airesintroduced, Corrientesintroduced, Córdobaintroduced, Entre Ríosintroduced, Formosaintroduced, La Pampaintroduced, Misionesintroduced); Argentina Northwest (Catamarcaintroduced, Jujuyintroduced, La Riojaintroduced, Mendozaintroduced, Saltaintroduced, San Juanintroduced, San Luisintroduced, Santiago del Esterointroduced, Tucumanintroduced); Argentina South (Chubutintroduced, Neuquénintroduced, Rio Negrointroduced, Santa Cruzintroduced, Santa Féintroduced, Tierra del Fuego (Argentina)introduced); Bahamas (Bahamasintroduced); Brazil South (Rio Grande do Sulintroduced, Santa Catarinaintroduced); Cayman Is. (Cayman Is.introduced); Chile Centralnative (Biobíointroduced, Coquimbointroduced, La Araucaniaintroduced, Mauleintroduced, O'Higginsintroduced, Santiagonative, Valparaísointroduced); Chile North (Antofagastaintroduced, Atacamaintroduced); Chile South (Aisénintroduced, Los Lagosintroduced, Magellanesnative); Colombia (Colombiaintroduced); Cuba (Cubaintroduced); Dominican Republic (Dominican Republicnative); Ecuadorintroduced; Guatemala (Guatemalaintroduced); Haiti (Haitinative, Navassa I.introduced); Hondurasintroduced; Jamaica (Jamaicaintroduced); Juan Fernández Is. (Juan Fernández Is.introduced); Leeward Is. (Virgin Is.introduced); Netherlands Antillesintroduced; Paraguay (Paraguayintroduced); Peruintroduced (Perunative); Puerto Rico (Puerto Ricointroduced); Uruguay (Uruguayintroduced); Venezuela (Venezuelanative); Windward Is. (Barbadosintroduced, Dominicaintroduced)

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 analysis in Kilian & al. (2009) 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 and unpubl.) 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) 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 is also a member of the Dubyaea-Nabalus-Soroseris-Syncalathium clade (Kilian & al. 2017).

References


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

Deng T., Zhang J.-W., Zhu X.-X., Zhang D.-G., Nie Z.-L. & Sun H. 2014: Youngia zhengyiana (Asteraceae, Crepidinae), a new species from south China, with notes on the systematics of Youngia inferred from morphology and nrITS phylogeny. – Phytotaxa 170: 259–268. // ➪ //

Enke N. & Gemeinholzer B. 2008: Babcock revisited: new insights into generic delimitation and character evolution in Crepis L. (Compositae: Cichorieae) from ITS and matK sequence data. – Taxon 57: 756-768.

Enke N., Kilian N., Nemomissa S. & Gemeinholzer G. 2008: Afroalpine Dianthoseris actually a congener of Crepis s.str. (Compositae, Cichorieae). – Bot. Jahrb. Syst. 127: 389-405.

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.

Kilian N., Sennikov A., Wang Z.-H., Gemeinholzer B. & Zhang J.-W. 2017[a]: Sub-Paratethyan origin and Middle to Late Miocene principal diversification of the Lactucinae (Cichorieae, Compositae) inferred from molecular phylogenetics, divergence-dating and biogeographic analysis. – Taxon 66: 675-703. // ➪ //

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Liu Y., Chen Y.-S. & Yang Q.-E. 2013: Generic status, circumscription, and allopolyploid origin of Faberia (Asteraceae: Cichorieae) as revealed by ITS and chloroplast DNA sequence data. – Taxon 62: 1235–1247. // ➪ //

Nakamura K., Chung S.-W., Kono Y., Ho M.-J., Hsu T.-C. & Peng C.-I 2014: Ixeridium calcicola (Compositae), a new limestone endemic from Taiwan, with notes on its atypical basic chromosome number, phylogenetic affinities, and a limestone refugium hypothesis. \u2013 PLoS ONE 9(10): e109797. // ➪ //

Pak J.-H. & Bremer K. 1995: Phylogeny and reclassification of the genus Lapsana (Asteraceae: Lactuceae). – Taxon 44: 13-21.

Pak J.-H. & Kawano S. 1992: Biosystematic studies on the genus Ixeris and its allied genera (Compositae: Lactuceae) 4. Taxonomic treatments and nomenclature. – Mem. Fac. Sci. Kyoto Univ., Ser. Biol. 15: 29-61.

Peng Y.-L., Zhang Y., Gao X.-F., Tong L.-J, Li L., Li R.-Y., Zhu Z.-M. & Xian J.-R. 2014: A phylogenetic analysis and new delimitation of Crepidiastrum (Asteraceae, tribe Cichorieae). – Phytotaxa 159: 241–255. // ➪ //

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Stebbins G. L. 1940: Studies in Cichorieae: Dubyaea and Soroseris. Endemics of the Sino-Himalayan region. – Mem. Torrey Bot. Club 19(3): 1-76.

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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.

Zhang J.-W., Sun H. & Nie E.-L. 2007: Karyological studies on the Sino-Himalayan endemic Soroseris and two related genera of tribe Lactuceae (Asteraceae). – Bot. J. Linn. Soc. 154: 79–87.

Zhang J. W., Nie Z. L. & Sun H. 2009: Cytological study on the genus Syncalathium (Asteraceae-Lactuceae), an endemic taxon to alpine scree of the Sino-Himalayas. – J. Syst. Evol. 47: 226–230.

Zhang J.-W., Nie Z. N., Wen J. & Sun H. 2011: Molecular phylogeny and biogeography of three closely related genera, Soroseris, Stebbinsia, and Syncalathium (Asteraceae, Cichorieae), endemic to the Tibetan Plateau, SW China. – Taxon 60: 15–26. // ➪ //

Credits

Zhang Jianwen 2009: Systematics (data).