Hypochaeridinae
Content
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 a sister clade (with Avellara, see below) to the remainder of 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
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 a sister clade (with Avellara, see below) to the remainder of 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
Distribution
Africa: Algeria native; Azores native; Canary Is. native; Cape Provinces (Eastern Cape Province introduced, Western Cape Province introduced); Cape Verde introduced; Chad native; Egypt native; Eritrea native; Ethiopia native; Free State introduced; Kenya introduced; KwaZulu-Natal introduced; Lesotho introduced; Libya native; Madagascar introduced; Madeira native; Mali native; Morocco native (Morocco native); Mozambique introduced; Nigeria native; Northern Provinces (Gauteng native, Mpumalanga introduced, North-West Province introduced, Northern Province introduced); Réunion introduced; Senegal native; Sudan native; Swaziland introduced; Tanzania introduced; Tunisia native; Zimbabwe introduced. Asia-Temperate: Afghanistan native; Altay native; Amur native; Buryatiya native; China North-Central (Beijing reported in error, Gansu native, Hebei native, Shaanxi native, Shandong native, Shanxi native); China South-Central (Guizhou native, Hubei native, Sichuan native, Yunnan native); China Southeast (Anhui native, Guangxi native, Henan native); Chita native; Cyprus native; East Aegean Is. native; Gulf States (Qatar native, United Arab Emirates native); Inner Mongolia (Nei Mongol native); Iran native; Iraq native; Irkutsk native; Japan (Hokkaido native, Honshu native, Kyushu native, Shikoku native); Kamchatka native; Kazakhstan native; Khabarovsk native; Kirgizistan native; Korea native (North Korea native, South Korea native); Krasnoyarsk native; Kuril Is. native; Kuwait native; Lebanon-Syria (Lebanon native, Syria native); Manchuria (Heilongjiang native, Jilin native, Liaoning native); Mongolia native ‒ native; Nansei-shoto introduced; North Caucasus native (Chechnya native, Dagestan native, Kabardino-Balkariya native, Karacheyevo-Cherkessiya native, Krasnodar native, Severo-Osetiya native, Stavropol native); Oman native; Palestine (Israel native, Jordan native); Primorye native; Qinghai native; Sakhalin native; Saudi Arabia native; Sinai native; Tadzhikistan native; Taiwan native; Tibet native; Transcaucasus (Abkhaziya native, Adzhariya native, Armenia native, Azerbaijan native, Georgia native, Nakhichevan native); Turkey native; Turkmenistan native; Tuva native; Uzbekistan native; West Siberia native; Xinjiang native; Yemen (North Yemen native, South Yemen native). Asia-Tropical: Assam (Assam introduced, Meghalaya native); Borneo (Sabah introduced); East Himalaya (Bhutan native, Sikkim native); India (Kerala introduced, Tamil Nadu introduced, Uttar Pradesh native); Myanmar native; Nepal native; Pakistan native; Sri Lanka introduced; Vietnam native; West Himalaya (Himachal Pradesh native, Jammu-Kashmir native). Australasia: Antipodean Is. introduced; Chatham Is. introduced; Kermadec Is. introduced; New South Wales (Australian Capital Territory native, New South Wales native); New Zealand North native; New Zealand South introduced; Northern Territory introduced; Queensland (Queensland native); South Australia native; Tasmania native; Victoria native; Western Australia (Western Australia native). Europe: Albania native ‒ 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 introduced (Czech Republic native, Slovakia native); Denmark native; East European Russia native; Finland native; France native (France native); Føroyar native; Germany native; Great Britain native; Greece native ‒ native; Hungary native; Iceland introduced; 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 ‒ native; Romania native; Sardegna native; Sicily native (Malta native, Sicily native); South European Russia native; Spain native (Andorra 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: Alabama introduced; Alaska introduced; Alberta introduced; Arizona introduced; Arkansas introduced; British Columbia introduced; California introduced; Colorado introduced; Connecticut introduced; Delaware introduced; District of Columbia introduced; Florida introduced; Georgia, U.S.A. introduced; Greenland introduced; Idaho introduced; Illinois introduced; Indiana introduced; Iowa introduced; Kansas introduced; Kentucky introduced; Labrador introduced; Louisiana introduced; Maine introduced; Maryland introduced; Massachusetts introduced; Mexican Pacific Is. (Guadalupe I. introduced); Mexico Central (México Distrito Federal introduced, México State introduced, Puebla introduced); Mexico Northeast (Coahuila introduced, Guanajuato introduced, Querétaro native); Mexico Northwest (Baja California introduced); Mexico Southwest (Michoacán introduced); Michigan introduced; Mississippi introduced; Missouri introduced; Montana introduced; Nevada introduced; New Brunswick introduced; New Hampshire introduced; New Jersey introduced; New Mexico introduced; New York introduced; Newfoundland (Newfoundland introduced, St.Pierre-Miquelon introduced); North Carolina introduced; North Dakota introduced; Northwest Territories introduced; Nova Scotia introduced; Ohio introduced; Oklahoma introduced; Ontario introduced; Oregon introduced; Pennsylvania introduced; Prince Edward I. introduced; Québec introduced; Rhode I. introduced ‒ introduced; Saskatchewan introduced; South Carolina introduced; Tennessee introduced; Texas introduced; Utah introduced; Vermont introduced; Virginia introduced; Washington introduced; West Virginia introduced; Wisconsin introduced. Pacific: Hawaii (Hawaiian Is. introduced); New Caledonia introduced. 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); Bolivia 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); 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 introduced; Dominican Republic introduced; Ecuador native; Guatemala introduced; Haiti (Haiti introduced); Jamaica introduced; Juan Fernández Is. introduced; Panamá introduced; Paraguay native; Peru native ‒ native; Uruguay native; Venezuela native
Bibliography
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