Hypochaeridinae

Primary tabs

Distribution

Africa: Algeria (Algerianative); Azores (Açôresnative); Canary Is. (Canary Is.native); Cape Provinces (Eastern Cape Provinceintroduced, Western Cape Provinceintroduced); Egypt (Egyptnative); Eritrea (Eritreanative); Ethiopia (Ethiopianative); Free State (Free Stateintroduced); Kenya (Kenyanative); KwaZulu-Natal (KwaZulu-Natalnative); Lesotho (Lesothointroduced); Libya (Libyanative); Madagascar (Madagascarintroduced); Madeira (Madeiranative); Mali (Malinative); Morocconative; Mozambique (Mozambiqueintroduced); Nigeria (Nigerianative); Northern Provinces (Gautengnative, Mpumalangaintroduced, North-West Provinceintroduced, Northern Provinceintroduced); Rwanda (Rwandanative); Réunion (Réunionintroduced); Senegal (Senegalnative); Swaziland (Swazilandintroduced); Tanzania (Tanzaniaintroduced); Tunisia (Tunisianative); Zaire (Zairenative); Zimbabwe (Zimbabweintroduced) Asia-Temperate: Afghanistan (Afghanistannative); Altay (Altaynative); Amur (Amurnative); Buryatiya (Buryatiyanative); China North-Central (Beijingreported in error, Gansunative, Hebeinative, Shaanxinative, Shandongnative, Shanxinative); China South-Central (Guizhounative, Hubeinative, Sichuannative, Yunnannative); China Southeast (Anhuinative, Guangxinative, Henannative); Chita (Chitanative); Cyprus (Cyprusnative); East Aegean Is. (East Aegean Is.native); Gulf States (Qatarnative, United Arab Emiratesnative); Inner Mongolia (Nei Mongolnative); Iran (Irannative); Iraq (Iraqnative); Irkutsk (Irkutsknative); Japan (Hokkaidonative, Honshunative, Kyushunative, Shikokunative); Kamchatka (Kamchatkanative); Kazakhstan (Kazakhstannative); Khabarovsk (Khabarovsknative); Kirgizistan (Kirgizistannative); Korea (North Koreanative, South Koreanative); Krasnoyarsk (Krasnoyarsknative); Kuril Is. (Kuril Is.native); Kuwait (Kuwaitnative); Lebanon-Syria (Lebanonnative, Syrianative); Manchuria (Heilongjiangnative, Jilinnative, Liaoningnative); Mongolianative (Mongolianative); Nansei-shoto (Nansei-shotointroduced); North Caucasusnative (Chechnyanative, Dagestannative, Kabardino-Balkariyanative, Karacheyevo-Cherkessiyanative, Krasnodarnative, Severo-Osetiyanative, Stavropolnative); Oman (Omannative); Palestine (Israelnative, Jordannative); Primorye (Primoryenative); Qinghai (Qinghainative); Sakhalin (Sakhalinnative); Saudi Arabia (Saudi Arabianative); Sinai (Sinainative); Tadzhikistan (Tadzhikistannative); Taiwanintroduced (Taiwannative); Tibet (Tibetnative); Transcaucasus (Abkhaziyanative, Adzhariyanative, Armenianative, Azerbaijannative, Georgianative, Nakhichevannative); Turkey (Turkeynative); Turkmenistan (Turkmenistannative); Tuva (Tuvanative); West Siberia (West Siberianative); Xinjiang (Xinjiangnative); Yemen (North Yemennative, South Yemennative) Asia-Tropical: Assam (Assamintroduced, Meghalayanative); Borneo (Sabahintroduced); East Himalaya (Bhutannative, Sikkimnative); India (Keralaintroduced, Tamil Naduintroduced, Uttar Pradeshnative); Myanmar (Myanmarnative); Nepal (Nepalnative); Pakistan (Pakistannative); Sri Lanka (Sri Lankaintroduced); Sumatera (Sumateranative); Vietnam (Vietnamnative); West Himalaya (Himachal Pradeshnative, Jammu-Kashmirnative) Australasia: Antipodean Is. (Antipodean Is.introduced); Chatham Is. (Chatham Is.introduced); Kermadec Is. (Kermadec Is.introduced); New South Wales (Australian Capital Territorynative, New South Walesnative); New Zealand North (New Zealand Northnative); New Zealand South (New Zealand Southintroduced); Northern Territory (Northern Territoryintroduced); Queensland (Queenslandnative); South Australia (South Australianative); Tasmania (Tasmanianative); Victoria (Victorianative); Western Australia (Western Australianative) 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; Føroyar (Føroyarnative); Germany (Germanynative); Great Britain (Great Britainnative); Greece (Greecenative); Hungary (Hungarynative); Iceland (Icelandintroduced); 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); Sweden (Swedennative); Switzerland (Switzerlandnative); Turkey-in-Europe (Turkey-in-Europenative); Ukraine (Moldovanative, Ukrainenative); Yugoslavia (Bosnia-Herzegovinanative, Croatianative, Macedonianative, Montenegronative, Serbianative, Slovenianative) Northern America: Alabama (Alabamaintroduced); Alaska (Alaskaintroduced); Alberta (Albertaintroduced); Arizona (Arizonaintroduced); Arkansas (Arkansasintroduced); British Columbia (British Columbiaintroduced); California (Californiaintroduced); Colorado (Coloradointroduced); Connecticut (Connecticutintroduced); Delaware (Delawarenative); District of Columbia (District of Columbiaintroduced); Florida (Floridanative); Georgia, U.S.A. (Georgia, U.S.A.native); Greenland (Greenlandintroduced); Idaho (Idahointroduced); Illinois (Illinoisintroduced); Indiana (Indianaintroduced); Iowa (Iowaintroduced); Kansas (Kansasintroduced); Kentucky (Kentuckyintroduced); Labrador (Labradorintroduced); Louisiana (Louisianaintroduced); Maine (Maineintroduced); Maryland (Marylandnative); Massachusetts (Massachusettsintroduced); Mexican Pacific Is. (Guadalupe I.introduced); Mexico Central (Mexico Distrito Federalintroduced, México Stateintroduced, Pueblaintroduced); Mexico Northeast (Coahuilaintroduced, Guanajuatointroduced, Querétaronative); Mexico Northwest (Baja Californiaintroduced); Mexico Southwest (Michoacánintroduced); Michigan (Michiganintroduced); Mississippi (Mississippinative); Missouri (Missouriintroduced); Montana (Montanaintroduced); Nevada (Nevadaintroduced); New Brunswick (New Brunswickintroduced); New Hampshire (New Hampshireintroduced); New Jersey (New Jerseynative); New Mexico (New Mexicointroduced); New York (New Yorkintroduced); Newfoundland (Newfoundlandintroduced, St.Pierre-Miquelonintroduced: adventitious (casual)); North Carolina (North Carolinanative); North Dakota (North Dakotaintroduced); Northwest Territories (Northwest Territoriesintroduced); Nova Scotia (Nova Scotiaintroduced); Ohio (Ohiointroduced); Oklahoma (Oklahomareported in error); Ontario (Ontariointroduced); Oregon (Oregonintroduced); Pennsylvania (Pennsylvaniaintroduced); Prince Edward I. (Prince Edward I.introduced); Québec (Québecintroduced); Rhode I.introduced (Rhode I.introduced); Saskatchewan (Saskatchewanintroduced); South Carolina (South Carolinanative); Tennessee (Tennesseeintroduced); Texas (Texasnative); Utah (Utahintroduced); Vermont (Vermontintroduced); Virginia (Virginianative); Washington (Washingtonintroduced); West Virginia (West Virginiaintroduced); Wisconsin (Wisconsinintroduced) Pacific: New Caledonia (New Caledoniaintroduced) Southern America: Argentina Northeast (Argentina Distrito Federalnative, Buenos Airesnative, Chaconative, Corrientesnative, Córdobanative, Entre Ríosnative, Formosanative, La Pampanative, Misionesnative); Argentina Northwest (Catamarcanative, Jujuynative, La Riojanative, Mendozanative, Saltanative, San Juannative, San Luisnative, Santiago del Esteronative, Tucumannative); Argentina South (Chubutnative, Neuquénnative, Rio Negronative, Santa Cruznative, Santa Fénative, Tierra del Fuego (Argentina)native); Bolivia (Bolivianative); Brazil South (Paranánative, Rio Grande do Sulnative, Santa Catarinanative); Brazil Southeast (Minas Geraisnative, Rio de Janeironative, São Paulonative); Chile Central (Biobíonative, Coquimbonative, La Araucanianative, Maulenative, O'Higginsnative, Santiagonative, Valparaísonative); Chile North (Antofagastanative, Atacamanative, Tarapacanative); Chile South (Aisénnative, Los Lagosnative, Magellanesnative); Colombia (Colombianative); Costa Rica (Costa Ricaintroduced); Dominican Republic (Dominican Republicintroduced); Ecuador (Ecuadornative); Guatemala (Guatemalaintroduced); Haiti (Haitinative); Jamaica (Jamaicaintroduced); Juan Fernández Is. (Juan Fernández Is.introduced); Paraguay (Paraguaynative); Peru (Perunative); Uruguayintroduced (Uruguaynative); Venezuela (Venezuelanative)

Systematics

The core of subtribe Hypochaeridinae is the Hypochaeris-Leontodon-Picris alliance. Tremetsberger & al. (2012) calculated a minimum age for the core Hypochaeridinae of c. 10.5 mya (95 % HPD interval = c. 7.5–13.5). Its relationship has been reconstructed by Samuel & al. (2003, 2006) and Enke & al. (2012) and includes 6 or 7 genera:
Hypochaeris, with a remarkably disjunct distribution of c. 15 species in the Mediterranean and Europe extending into NE Asia and 40-45 species in South America (Tremetsberger & al. 2005), has colonised South America apparently via long-distant dispersal from the W Mediterranean (Cerbah & al. 1999; Samuel & al. 2003; Weiss-Schneeweiss & al. 2003, 2008; Tremetsberger & al. 2005).
The central Mediterranean monospecifc Robertia, formerly included in Hypochaeris, had been questioned as a member of the latter genus by Samuel & al. (2003) and its status as a separate lineage has been corroborated by Enke & al. (2012).
Picris and Helminthotheca have been confirmed as separate monophyletic lineages in the nrITS phylogenies by Samuel & al. (2006) and Enke & al. (2012).
Leontodon in its previous circumscription was clearly revealed as diphyletic by Samuel & al. (2006), necessitating the recognition of L. subg. Oporinia as separate genus Scorzoneroides (Greuter & al. 2006). The results of the molecular phylogenetic analyses by Samuel & al. (2006) and Enke & al. (2012) moreover raise the question whether Leontodon in the narrower sense excluding Scorzoneroides is monophyletic with respect to L. sect. Thrincia. While the matK plastid DNA phylogeny by Samuel & al. (2006) is inconclusive due to lacking resolution, L. sect. Thrincia forms a statistically well supported lineage separate from the remainder of Leontodon s.str. (L. sect. Leontodon and Astherotrix) in the nrITS phylogenies by Samuel & al. (2006) and Enke & al. (2012). Considering that L. sect. Thrincia is also morphologically well distinct from the remainder of Leontodon s.str., Talavera & al. (2015) and Talavera & Talavera (2017) resurrected the former genus Thrincia for the eight western Mediterranean centred species included. This taxonomic conclusion is plausible but not inevitable from the molecular phylogenetic analyses currently at hand. The deeper nodes of the Picris-Helmintotheca-Leontodon-Hedypnois-Thrincia clade are not resolved in the nrITS phylogeny by Samuel & al. (2006). The grouping of L. sect. Thrincia in a trichotomy with Picris and Helminthotheca in the nrITS tree by Enke & al. (2012) has no statistical support. The tree based on the combined matK-nrITS dataset by Samuel & al. (2006), however, resolves a moderately supported sister group relationship between L. sect. Thrincia and the remainder of Leontodon. The molecular phylogenetic results by Moura & al. (2015) are inconclusive with respect to the position of L. sect. Thrincia due to their restricted sampling. Since there is no conclusive evidence against a sistergroup relationship of L. sect. Thrincia and the remainder of Leontodon s.str., the Cichorieae Systematics Portal for the time being maintains the wider circumscription of Leontodon with inclusion of L. sect. Thrincia.
Hedypnois, the only genus in this alliance with a scabrid instead of a plumose pappus, indicates a reversal from the plumose pappus back to a scabrid or scaly pappus. This genus has been shown, however, to be nested in Leontodon by Enke & al. (2012); if paraphylectic genera are not accepted, it has thus to be considered a congener of Leontodon.

The placement of Urospermum in the Hypochaeridinae has been confirmed in the recent molecular phylogenetic analysis by Fernández-Mazuecos & al. (2016). This bispecific, predominantly Mediterranean genus with plumose Hypochaeris type pappus, a hispid indumentum, and a pollen type very similar to Hypochaeris and Picris (Lack & Leuenberger 1979) has in spite of its uniseriate, basally connate involucre and unique achenes unequivocally been placed into the Hypochaeridinae since Hoffmann (1890-94).

The genus Avellara, with its single species A. fistulosa, a rare, critically endangered endemic of the Iberian Peninsula was originally placed in Scorzonera but has to be excluded from this genus as well as from the Scorzonerinae for morphological, palynological and cytological evidence as was shown by Blanca & Díaz (1985), and for molecular phylogenetic evidence provided by Fernández-Mazuecos & al. (2016). It apparently represents an isolated, early diverging lineage of the large Chondrillinae-Crepidinae-Hyoserdinae-Hypochaeridinae-Lactucinae clade of the Cichorieae with affinities to Urospermum. The divergence of Avellara and Urospermum is estimated to have taken place already in the Middle to Late Miocene, around 15.58–8.62 mya, thus in a time where the lineages of the major Cichorieae clades diversified. For the time being, this genus is treated as a member of the Hypochaeridinae.

In contrast to the circumscription of the Hypochaeridinae by Bremer (1994) and Lack (2006) several genera have been excluded in the revised circumscription based on recent molecular phylogenetic analyses by Gemeinholzer & al. (in Kilian & al. 2009): (1) Aposeris and Hyoseris group with the Sonchus-Launaea-Reichardia alliance (Hyoseridinae), (2) Rhagadiolus (confirming an earlier finding by Whitton & al. 1995) and Garhadiolus are nested in subtribe Crepidinae and (3) the relationship of Arnoseris to the Tolpis alliance (Cichoriinae) rather than to the Hypochaeris alliance has been confirmed.

In various nrITS trees published (Koopman & al. 1998; Kilian & al. 2009; Enke & al. 2012; Wang & al. 2013, Fernández-Mazuecos & al. 2016) Prenanthes purpurea (providing the type of the name Prenanthes) was found nested in the Hypochaeridinae clade. It appeared there as sister to the core Hypochaeridinae with Urospermum as the sister-group to the latter two (Kilian & al. 2009; Enke & al. 2012). This result remained enigmatic because morphologically as well as cytologically P. purpurea and the Hypochaeridinae are entirely unrelated, while a relationship with subtribe Lactucinae as revealed in plastid DNA marker trees is much more plausible (Wang & al. 2013). Kilian & al. (2017) finally have shown that the grouping of P. purpurea with the Hypochaeridinae is apparently the result of insufficient sampling regarding early diverging lineages and long branch attraction. In their more densely sampled nrITS tree P. purpurea is resolved with statistical support as an early diverging lineage of the Lactucinae.A,B,C,D,E,F,G,H,I,J,K,L,M,N,O,P,Q,R,S,T,U,V,W

Bibliography

A. Blanca G. & Díaz de la Guardia C., Avellara, género nuevo de la tribu Lactuceae (Compositae) en la península Ibérica in Candollea 40: 447-458. 1985
B. Bremer K., Asteraceae. Cladistics and classification. 1994
C. Cerbah M., Coulaud J., Brown S. C. & al., Evolutionary DNA variation in the genus Hypochaeris (Asteraceae) in Heredity 82: 261-266. 1999
D. Enke N., Gemeinholzer B. & Zidorn C., Molecular and phytochemical systematics of the subtribe Hypochaeridinae (Asteraceae, Cichorieae) in Organisms Diversity Evol. 12: 1-16. 2012
E. Fernández-Mazuecos M. & al., Narrow endemics on coastal plains: Miocene divergence of the critically endangered genus Avellara (Compositae) in Pl. Biol. 18: 729-738. 2016
F. Greuter W., Gutermann W. & Talavera S., A preliminary conspectus of Scorzoneroides (Compositae, Cichorieae) with validation of the required new names in Willdenowia 36: 689–692. 2006
G. Hoffmann O. - Compositae in Engler A. & Prantl K., Die natürlichen Pflanzenfamilien 4(5). 1890-1894
H. Kilian N., Gemeinholzer B. & Lack H. W. - Tribe Cichorieae in Funk V. A., Susanna A., Stuessy T. & al., Systematics, evolution and biogeography of Compositae. 2009
I. Kilian N., Sennikov A. N., Wang Z. H. & al., Sub-Paratethyan origin and Middle to Late Miocene principal diversification of the Lactucinae (Cichorieae, Compositae) inferred from molecular phylogenetics, divergence-dating and biogeographic analysis in Taxon 66: 675-703. 2017
J. Koopman W. J., Guetta E., Wiel C. C. M. van de & al., Phylogenetic relationships among Lactuca (Asteraceae) species and related genera based on ITS-1 DNA sequences in Amer. J. Bot. 85: 1517–1530. 1998
K. Lack H. W. & Leuenberger B., Pollen and taxonomy of Urospermum (Asteraceae, Lactuceae) in Pollen Spores 21: 415–425. 1979
L. Lack H. W. - Cichorieae in Kadereit J. W. & Jeffrey C., The families and genera of vascular plants 8. 2006
M. Moura M. & al., A revision of the genus Leontodon (Asteraceae) in the Azores based on morphological and molecular evidence in Phytotaxa 210. 2015
N. Samuel R., Gutermann W., Stuessy T. & al., Molecular phylogenetics reveals Leontodon (Asteraceae, Cichorieae) to be diphyletic in Amer. J. Bot. 93: 1193-1205. 2006
O. Samuel R., Stuessy T., Tremetsberger K. & al., Phylogenetic relationships among species of Hypochaeris (Asteraceae, Cichorieae) based on ITS, plastid trnL intron, trnL-F spacer, and matK sequences in Amer. J. Bot. 90: 496–507. 2003
P. Talavera S. & Talavera M. - Thrincia in Talavera S. & al. (ed.), Flora iberica 16(2). Compositae (partim). 2017
Q. Talavera S., Talavera M. & Sánchez C., Los géneros Thrincia Roth y Leontodon L. (Compositae, Cichorieae) en Flora Iberica in Acta Bot. Malac. 40: 344-364. 2015
R. Tremetsberger K., Gemeinholzer B., Zetzsche H. & al., Divergence time estimation in Cichorieae (Asteraceae) using a fossil-calibrated relaxed molecular clock in Organisms Diversity Evol. 13: 1-13. 2012
S. Tremetsberger K., Weiss-Schneeweiss H., Stuessy T. & al., Nuclear ribosomal DNA and karyotypes indicate a NW African origin of South American Hypochaeris (Asteraceae, Cichorieae) in Molec. Phylogen. Evol. 35: 102-116. 2005
T. Wang Z. H., Peng H. & Kilian N., Molecular phylogeny of the Lactuca alliance (Cichorieae subtribe Lactucinae, Asteraceae) with focus on their Chinese centre of diversity detects potential events of reticulation and chloroplast capture in PLoS One 8(12), e82692. 2013
U. Weiss-Schneeweiss H., Stuessy T., Siljak-Yakovlev S. & al., Karyotype evolution in South American species of Hypochaeris (Asteraceae, Lactuceae) in Pl. Syst. Evol. 241: 171-184. 2003
V. Weiss-Schneeweiss H., Tremetsberger K., Schneeweiss G. M. & al., Karyotype diversification and evolution in diploid and polyploid South American Hypochaeris (Asteraceae) inferred from rDNA localization and genetic fingerprint data in Ann. Bot. (Oxford) 101: 909-918. 2008
W. Whitton J., Wallace R. S. & Jansen R. K, Phylogenetic relationships and patterns of character change in the tribe Lactuceae (Asteraceae) based on chloroplast DNA restriction site variation in Canad. J. Bot. 73: 1058-1073. 1995