Hyoseridinae
Systematics
Extensive molecular phylogenetic analyses of the Sonchus alliance by Kim & al. (1996, 1997, 1999a,b, 2004, 2007) revealed that the genus Dendroseris, and, by morphology, also the monospecific genus Thamnoseris, both formerly included in a separate subtribe Dendroseridinae, are part of the Sonchinae and the Sonchus alliance in particular.
Recent molecular phylogenetic analyses by Gemeinholzer & al. (in Kilian & al. 2009) revealed that also Aposeris (lacking a pappus) and Hyoseris (with an inner pappus of basally strongly widened bristles) have to be included in the Sonchinae. Blackmore (1981) stated that the palynological evidence is inconclusive for the placement of Hyoseris, reflecting a possible relationship to the Hypochaeris alliance as likely as to the Launaea-Reichardia-Sonchus alliance. Molecular analyses by Samuel & al. (2003) revealed that neither species is a member of the Hypochaeridinae. Aposeris is sister to all other genera of this subtribe, which are rather closely related to each other, while Hyoseris is sister to the clade including Launaea, Reichardia and the Sonchus alliance. Inclusion of Hyoseris required a change of name of the subtribe from Sonchinae to Hyoseridinae because of the priority rule.
The various species-poor genera, established within the Sonchus alliance for species of the Canary Islands (Chrysoprenanthes, Babcockia, Lactucosonchus, Sventenia, Taeckholmia, Wildpretia) and of Australia/New Zealand (Actites, Embergia, Kirkianella), were nested within Sonchus in all recent molecular phylogenetic analyses (Kim & al. 1996 [a, b], 1999 [a, b], 2004, 2007). The same has been confirmed for the monospecific Mediterranean Aetheorhiza as well as for Dendroseris and Thamnoseris, endemic to the Pacific Juan Fernandez and Desventuradas Islands, respectively (Kim & al. 2007). The Canarian, Mediterranean and Australian/New Zealandean genera are morphologically questionably distinct from Sonchus and their inclusion in Sonchus seems therefore a justified consequence. The case may appear different with respect to Dendroseris (Carlquist 1966; Crawford & al. 1992; Sang & al. 1994) and Thamnoseris, which apparently have evolved from within Sonchus after long-distant dispersal to these Pacific island groups. They may be exemplary for the need to accept, whenever appropriate, also paraphyletic genera as long as we want taxonomy to reflect evolution (Hörandl 2007, Brummitt 2008). Mejías & Kim (2012), however, came, also after an extensive review of the literature on these genera, to the conclusion that Dendroseris and Thamnoseris should best be included in Sonchus and provided the corresponding nomenclatural combinations. The treatment here follows these authors.A,B,C,D,E,F,G,H,I,J,K,L,M,N,O
Recent molecular phylogenetic analyses by Gemeinholzer & al. (in Kilian & al. 2009) revealed that also Aposeris (lacking a pappus) and Hyoseris (with an inner pappus of basally strongly widened bristles) have to be included in the Sonchinae. Blackmore (1981) stated that the palynological evidence is inconclusive for the placement of Hyoseris, reflecting a possible relationship to the Hypochaeris alliance as likely as to the Launaea-Reichardia-Sonchus alliance. Molecular analyses by Samuel & al. (2003) revealed that neither species is a member of the Hypochaeridinae. Aposeris is sister to all other genera of this subtribe, which are rather closely related to each other, while Hyoseris is sister to the clade including Launaea, Reichardia and the Sonchus alliance. Inclusion of Hyoseris required a change of name of the subtribe from Sonchinae to Hyoseridinae because of the priority rule.
The various species-poor genera, established within the Sonchus alliance for species of the Canary Islands (Chrysoprenanthes, Babcockia, Lactucosonchus, Sventenia, Taeckholmia, Wildpretia) and of Australia/New Zealand (Actites, Embergia, Kirkianella), were nested within Sonchus in all recent molecular phylogenetic analyses (Kim & al. 1996 [a, b], 1999 [a, b], 2004, 2007). The same has been confirmed for the monospecific Mediterranean Aetheorhiza as well as for Dendroseris and Thamnoseris, endemic to the Pacific Juan Fernandez and Desventuradas Islands, respectively (Kim & al. 2007). The Canarian, Mediterranean and Australian/New Zealandean genera are morphologically questionably distinct from Sonchus and their inclusion in Sonchus seems therefore a justified consequence. The case may appear different with respect to Dendroseris (Carlquist 1966; Crawford & al. 1992; Sang & al. 1994) and Thamnoseris, which apparently have evolved from within Sonchus after long-distant dispersal to these Pacific island groups. They may be exemplary for the need to accept, whenever appropriate, also paraphyletic genera as long as we want taxonomy to reflect evolution (Hörandl 2007, Brummitt 2008). Mejías & Kim (2012), however, came, also after an extensive review of the literature on these genera, to the conclusion that Dendroseris and Thamnoseris should best be included in Sonchus and provided the corresponding nomenclatural combinations. The treatment here follows these authors.A,B,C,D,E,F,G,H,I,J,K,L,M,N,O
Distribution
Africa: Aldabra native; Algeria native; Angola native; Azores native; Benin native; Burkina native; Burundi native; Cameroon native; Canary Is. native; Cape Provinces (Eastern Cape Province native, Northern Cape Province native, Western Cape Province native); Cape Verde native ‒ native; Central African Republic native; Chad native; Comoros (Comoros native, Mayotte introduced); Congo native; Djibouti native; Egypt native; Equatorial Guinea native; Eritrea native; Ethiopia native; Free State native; Gabon native; Gambia, The introduced; Ghana native; Guinea native; Gulf of Guinea Is. (Bioko native, São Tomé introduced); Ivory Coast native; Kenya native; KwaZulu-Natal native; Lesotho native; Libya native; Madagascar native; Madeira native; Malawi native; Mali native; Mauritania native; Mauritius native; Morocco native (Morocco native); Mozambique native; Namibia native; Niger native; Nigeria native; Northern Provinces (Gauteng native, Mpumalanga native, North-West Province native, Northern Province native); Rwanda native; Réunion native; Selvagens native; Senegal native; Seychelles native; Sierra Leone native; Socotra native; Somalia native; St.Helena native; Sudan native; Swaziland native; Tanzania native; Togo native; Tunisia native; Uganda native; Western Sahara native ‒ native; Zaire native; Zambia native; Zimbabwe native. Asia-Temperate: Afghanistan native; Altay native; Amur native; Buryatiya native; China North-Central (Gansu native, Hebei native, Shaanxi native, Shandong native, Shanxi native); China South-Central (Guizhou native, Hubei native, Sichuan native, Yunnan native); China Southeast (Anhui introduced, Fujian native, Guangdong native, Guangxi native, Henan native, Hong Kong native, Hunan native, Jiangsu native, Jiangxi native, Zhejiang native); Chita native; Cyprus native; East Aegean Is. native; Gulf States (Bahrain native, Qatar native, United Arab Emirates native); Hainan native; Inner Mongolia (Nei Mongol native, Ningxia native); Iran native; Iraq native; Irkutsk native; Japan introduced (Hokkaido native, Honshu native, Kyushu native, Shikoku native); Kamchatka doubtfully present; Kazakhstan native; Khabarovsk native; Kirgizistan native; Korea native (South Korea native); Krasnoyarsk native; Kuril Is. native; Kuwait native; Lebanon-Syria (Lebanon native, Syria native); Magadan doubtfully present; Manchuria (Heilongjiang native, Jilin native, Liaoning native); Mongolia native; North Caucasus native (Chechnya native, Dagestan native, Kabardino-Balkariya native, Karacheyevo-Cherkessiya native, Krasnodar native, Severo-Osetiya native, Stavropol native); Ogasawara-shoto introduced; 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 doubtfully present; Uzbekistan native; West Siberia native; Xinjiang native; Yakutskiya native; Yemen (North Yemen native, South Yemen native). Asia-Tropical: Andaman Is. (Andaman Is. native); Assam (Assam native, Manipur native, Meghalaya native); Bangladesh native; Borneo (Sabah introduced); East Himalaya (Bhutan native, Sikkim native); India introduced (Bihar native, Chattisgarh native, Haryana native, Karnataka native, Kerala native, Madhya Pradesh native, Maharashtra native, Orissa native, Punjab native, Rajasthan native, Tamil Nadu native, Uttar Pradesh native, West Bengal native); Jawa native; Laos native; Malaya (Peninsular Malaysia native); Maldives native; Myanmar native; Nepal native; New Guinea reported in error (Irian Jaya introduced, Papua New Guinea introduced); Pakistan native ‒ native; Philippines native; Sri Lanka native; Sumatera native; Thailand native; Vietnam native; West Himalaya (Himachal Pradesh native, Jammu-Kashmir native). Australasia: Antipodean Is. introduced; 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 (Ashmore-Cartier Is. introduced, 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 (France native); Føroyar native; Germany native; Great Britain native; Greece native; Hungary native; Iceland introduced; Ireland native (Ireland native, Northern Ireland 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); 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; Manitoba introduced; Maryland introduced; Massachusetts introduced; Mexican Pacific Is. (Guadalupe I. introduced); Mexico Central (Morelos introduced, México Distrito Federal introduced, México State introduced, Puebla introduced, Tlaxcala introduced); Mexico Gulf (Veracruz introduced); Mexico Northeast (Chihuahua introduced, Coahuila introduced, Durango introduced, Guanajuato introduced, Hidalgo introduced, Nuevo León introduced, Querétaro introduced, San Luis Potosí introduced, Tamaulipas introduced, Zacatecas introduced); Mexico Northwest (Baja California introduced, Baja California Sur introduced, Sinaloa introduced, Sonora introduced); Mexico Southeast (Campeche introduced, Chiapas introduced, Quintana Roo introduced, Yucatán introduced); Mexico Southwest (Colima introduced, Guerrero introduced, Jalisco introduced, Michoacán introduced, Nayarit introduced, Oaxaca introduced); Michigan introduced; Minnesota introduced; Mississippi introduced; Missouri introduced; Montana introduced; Nebraska 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; Nunavut introduced; Ohio introduced; Oklahoma introduced; Ontario introduced; Oregon introduced; Pennsylvania introduced; Prince Edward I. introduced; Québec introduced; Rhode I. introduced; Saskatchewan introduced; South Carolina introduced; South Dakota introduced; Tennessee introduced; Texas introduced; Utah introduced; Vermont introduced; Virginia introduced; Washington introduced; West Virginia introduced; Wisconsin introduced; Wyoming introduced; Yukon introduced. Pacific: Cook Is. introduced; Fiji introduced; Hawaii (Hawaiian Is. introduced); New Caledonia introduced. Southern America: Argentina Northeast (Argentina Distrito Federal introduced, Buenos Aires introduced, Chaco introduced, Corrientes introduced, Córdoba introduced, Entre Ríos introduced, Formosa introduced, La Pampa introduced, Misiones introduced); Argentina Northwest (Catamarca introduced, Jujuy introduced, La Rioja introduced, Mendoza introduced, Salta introduced, San Juan introduced, San Luis introduced, Santiago del Estero introduced); Argentina South (Chubut introduced, Neuquén introduced, Rio Negro introduced, Santa Cruz introduced, Santa Fé introduced, Tierra del Fuego (Argentina) introduced); Aruba introduced; Bahamas introduced; Belize introduced; Bolivia introduced; Brazil Northeast (Bahia introduced); Brazil South (Paraná introduced, Rio Grande do Sul introduced, Santa Catarina introduced); Brazil Southeast (Trindade introduced); Cayman Is. introduced; Chile Central (Biobío introduced, Coquimbo introduced, La Araucania introduced, Maule introduced, O'Higgins introduced, Santiago introduced, Valparaíso introduced); Chile North (Antofagasta introduced, Atacama introduced, Tarapaca introduced); Chile South (Aisén introduced, Los Lagos introduced, Magellanes introduced); Colombia introduced; Costa Rica introduced; Cuba introduced; Desventurados Is. native; Dominican Republic introduced; Guatemala introduced; Haiti (Haiti introduced); Jamaica introduced; Juan Fernández Is. native ‒ native; Leeward Is. (Anguilla introduced, Antigua-Barbuda introduced, Guadeloupe introduced, Montserrat introduced, Netherlands Leeward Is. introduced, St.Martin-St.Barthélémy introduced, Virgin Is. introduced); Netherlands Antilles (Bonaire introduced, Curaçao introduced); Nicaragua introduced; Panamá introduced; Paraguay introduced; Peru introduced; Puerto Rico introduced; Trinidad-Tobago introduced; Turks-Caicos Is. introduced; Uruguay introduced; Venezuela introduced; Venezuelan Antilles introduced; Windward Is. (Barbados introduced, Dominica introduced, Grenada introduced, Martinique introduced, St.Lucia introduced, St.Vincent introduced)
Bibliography
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B. Blackmore S. 1981: Palynology and intergeneric relationships in subtribe Hyoseridinae (Compositae: Lactuceae). – Botanical Journal of the Linnean Society 82: 1-13
C. Brummitt R. K. 2008: Evolution in taxonomic perspective. – Taxon 57: 1049-1050. http://doi.org/10.1002/tax.574002
D. Carlquist S. 1966: The biota of long-distance dispersal II. Loss of dispersibility in Pacific Compositae. – Evolution 20: 30-48. http://doi.org/10.1111/j.1558-5646.1966.tb03341.x
E. Hörandl E. 2007: Neglecting evolution is bad taxonomy. – Taxon 56: 1-5. http://doi.org/10.2307/25065730
F. Kilian N., Gemeinholzer, B. & Lack H. W. 2009: Tribe Cichorieae, 343–383. – In: Funk V. A., Susanna A., Stuessy T. & Bayer R. J., Systematics, evolution and biogeography of Compositae. – Vienna: IAPT
G. Kim S.-C., Crawford D. J. & Jansen R. K 1996: Phylogenetic relationships among the genera of the subtribe Sonchinae (Asteraceae): evidence from ITS sequences. – Syst. Bot. 21: 417–432. http://doi.org/10.2307/2419668
H. Kim S.-C., Crawford D. J., Francisco-Ortega J. & Santos-Guerra A. 1996: A common origin for woody Sonchus and five related genera in the Macaronesian islands: molecular evidence for extensive radiation. – Proc. Natl. Acad. Sci. USA 93: 7743-7748. http://doi.org/10.1073/pnas.93.15.7743
I. Kim S.-C., Crawford D. J., Francisco-Ortega J. & Santos-Guerra A. 1999: Adaptive radiation and genetic differentiation in the woody Sonchus alliance (Asteraceae: Sonchinae) in the Canary Islands. – Pl. Syst. Evol. 215: 101-118. http://doi.org/10.1007/BF00984650
J. Kim S.-C., Crawford D. J., Jansen R. K & Santos-Guerra A. 1999: The use of a non-coding region of chloroplast DNA in phylogenetic studies of the subtribe Sonchinae (Asteraceae: Lactuceae). – Pl. Syst. Evol. 215: 85-99. http://doi.org/10.1007/BF00984649
K. Kim S.-C., Lee C. & Mejías J. A. 2007: Phylogenetic analysis of chloroplast DNA matK gene and ITS of nrDNA sequences reveals polyphyly of the genus Sonchus and new relationships among the subtribe Sonchinae (Asteraceae: Cichorieae). – Molec. Phylogen. Evol. 44: 578-597
L. Kim S.-C., Lu C. T. & Lepschi B. J. 2004: Phylogenetic positions of Actites megalocarpa and Sonchus hydrophilus (Sonchinae: Asteraceae) based on ITS and chloroplast non-coding DNA sequences. – Austral. Syst. Bot. 17: 73-81. http://doi.org/10.1071/SB03019
M. Mejías J. A. & Kim S.-C. 2012: Taxonomic treatment of Cichorieae (Asteraceae) endemic to the Juan Fernandez and Desventuradas Islands (SE Pacific). – Annales botanici fennici 49: 171-178. http://doi.org/10.5735/085.049.0303
N. Samuel R., Stuessy T., Tremetsberger K., Baeza C. M. & Siljak-Yakovlev S. 2003: Phylogenetic relationships among species of Hypochaeris (Asteraceae, Cichorieae) based on ITS, plastid trnL intron, trnL-F spacer, and matK sequences. – Amer. J. Bot. 90: 496–507
O. Sang T., Crawford D. J., Kim S.-C. & Stuessy T. 1994: Radiation of the endemic genus Dendroseris (Asteraceae) on the Juan Fernandez Islands: evidence from sequences of the ITS regions of nuclear ribosomal DNA. – Amer. J. Bot. 81: 1494-1501. http://doi.org/10.1002/j.1537-2197.1994.tb15635.x