Scolyminae
Content
Distribution
Africa: Algeria native; Azores introduced; Canary Is. native; Egypt native; Libya native; Madeira native; Morocco native; Sudan native; Tunisia native Asia-Temperate: Afghanistan doubtfully present; Cyprus native; East Aegean Is. native; Iran native ‒ native; Iraq native ‒ native; Kazakhstan doubtfully present; Lebanon-Syria (Lebanon native, Syria native); North Caucasus native (Krasnodar native); Palestine (Israel native, Jordan native); Sinai native; Transcaucasus (Abkhaziya native, Adzhariya native, Armenia native, Azerbaijan native, Georgia native, Nagorno Karabakh doubtfully present, Nakhichevan native); Turkey native ‒ native; Turkmenistan doubtfully present; Uzbekistan doubtfully present Australasia: New South Wales (New South Wales introduced); Northern Territory introduced; Queensland (Queensland introduced); South Australia introduced; Victoria introduced Europe: Albania native; Baleares native; Belgium (Belgium introduced: adventitious (casual)); Bulgaria native; Corse native; Czechoslovakia (Czech Republic introduced); France native; Greece native; Italy native; Kriti native; Krym native; Poland introduced: adventitious (casual); Portugal native; Romania native; Sardegna native; Sicily native (Malta native, Sicily native); Spain native (Andorra native, Gibraltar native, Spain native); Switzerland introduced; Turkey-in-Europe native; Ukraine (Ukraine native); Yugoslavia (Bosnia-Herzegovina native, Croatia native, Macedonia native: presence questionable, Montenegro native, Serbia native, Slovenia native) Northern America: Alabama introduced; California introduced; New York introduced; North Carolina introduced; Pennsylvania introduced Southern America: Argentina Northeast (Buenos Aires introduced, Entre Ríos introduced)
Systematics
The Scolyminae are treated here in the revised circumscription based on the molecular phylogenetic analyses by Gemeinholzer & al. (in Kilian & al. 2009) and Tremetsberger & al. (2012), comprising four genera with about a dozen species. According to these authors, within the Scolyminae, Catananche is sister to the clade including Hymenonema and Scolymus, while Gundelia is sister to all three genera. For the inclusion of Gundelia see also under Cichorieae: Systematics: Circumscription of the Cichorieae.
Usually Scolymus was treated as an isolated genus without apparent relationship and thus, if formally placed in a subtribe, as the only member of the Scolyminae (Jeffrey 1966; Bremer 1994; Lack 2006). Catananche and Hymenonema, on the other hand, were placed together with Rothmaleria in a separate subtribe (or informal entity, respectively, by Jeffrey (1966), Bremer (1994) and Lack (2006). As an exception, Blackmore (1981), however, for palynological reasons already associated Scolymus with Catananche and Hymenonema, but also saw Rothmaleria as a member of this alliance, which he treated as subtribe Scolyminae. According to the molecular analyses by Gemeinholzer & al. (in Kilian & al. 2009), Rothmaleria is not related to Catananche and Hymenonema but to Tolpis, as Stebbins (1953) already assumed from morphological data, and therefore placed in subtribe Cichoriinae.
Morphologically the Scolyminae appear to be a rather diverse subtribe. They are characterised by an annual to perennial life form, entire to pinnatisect or coarsely lobed, pinnatisect leaves, heads with receptacular scales or bristles, and the pappus being either absent or of scabrid bristles or lanceolate scales. Both Gundelia and Scolymus are spiny leafy herbs, the only ones in the entire tribe, with similar pollen morphology and also with sessile heads (or syncalathia, respectively). Moreover, they share the presence of both (functional) oil ducts and latex canals in the roots; functional oil ducts are very rare in the tribe, being otherwise present only in Scorzonera (see Cichorieae: Systematics). Gundelia morphologically holds an isolated position because of its heads with non-ligulate but tubular flowers and its much-derived synflorescence, in which one-flowered heads (with much reduced involucres) form a secondary head (syncalathium), of which again a few dozens are aggregated in a second order syncalathium (Classen-Bockhoff & al. 1989). Scolymus and Hymenonema share a chromosome number of 2n = 20, while Gundelia and Catananche have 2n = 18.
Blackmore S. 1981: Palynology and intergeneric relationships in subtribe Hyoseridinae (Compositae: Lactuceae). – Bot. J. Linn. Soc. 82: 1-13.
Bremer K. 1994: Asteraceae. Cladistics and classification. – Portland: Timber.
Classen-Bockhoff R., Froebe H. A. & Langerbeins D. 1989: Die Infloreszenzstruktur von Gundelia tournefortii L. (Asteraceae). – Flora 182: 463-479.
Jeffrey C. 1966: Notes in Compositae I. The Cichorieae in East Tropical Africa. – Kew Bulletin 18: 427-486.
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.
Lack H. W. 2006: Tribe Cichorieae Lam. & DC. – Pp. 180-199 in: Kadereit J. W. & Jeffrey C. (ed.), The families and genera of vascular plants 8. – Berlin: Springer.
Stebbins G. L. 1953: A new classification of the tribe Cichorieae, family Compositae. – Madroño 12: 65–81.
Tremetsberger K., Gemeinholzer B., Zetzsche H., Blackmore S., Kilian N. & Talavera S. 2012: Divergence time estimation in Cichorieae (Asteraceae) using a fossil-calibrated relaxed molecular clock. – Org. Divers. Evol. 13: 1-13. // ➪ //
Usually Scolymus was treated as an isolated genus without apparent relationship and thus, if formally placed in a subtribe, as the only member of the Scolyminae (Jeffrey 1966; Bremer 1994; Lack 2006). Catananche and Hymenonema, on the other hand, were placed together with Rothmaleria in a separate subtribe (or informal entity, respectively, by Jeffrey (1966), Bremer (1994) and Lack (2006). As an exception, Blackmore (1981), however, for palynological reasons already associated Scolymus with Catananche and Hymenonema, but also saw Rothmaleria as a member of this alliance, which he treated as subtribe Scolyminae. According to the molecular analyses by Gemeinholzer & al. (in Kilian & al. 2009), Rothmaleria is not related to Catananche and Hymenonema but to Tolpis, as Stebbins (1953) already assumed from morphological data, and therefore placed in subtribe Cichoriinae.
Morphologically the Scolyminae appear to be a rather diverse subtribe. They are characterised by an annual to perennial life form, entire to pinnatisect or coarsely lobed, pinnatisect leaves, heads with receptacular scales or bristles, and the pappus being either absent or of scabrid bristles or lanceolate scales. Both Gundelia and Scolymus are spiny leafy herbs, the only ones in the entire tribe, with similar pollen morphology and also with sessile heads (or syncalathia, respectively). Moreover, they share the presence of both (functional) oil ducts and latex canals in the roots; functional oil ducts are very rare in the tribe, being otherwise present only in Scorzonera (see Cichorieae: Systematics). Gundelia morphologically holds an isolated position because of its heads with non-ligulate but tubular flowers and its much-derived synflorescence, in which one-flowered heads (with much reduced involucres) form a secondary head (syncalathium), of which again a few dozens are aggregated in a second order syncalathium (Classen-Bockhoff & al. 1989). Scolymus and Hymenonema share a chromosome number of 2n = 20, while Gundelia and Catananche have 2n = 18.
References
Blackmore S. 1981: Palynology and intergeneric relationships in subtribe Hyoseridinae (Compositae: Lactuceae). – Bot. J. Linn. Soc. 82: 1-13.
Bremer K. 1994: Asteraceae. Cladistics and classification. – Portland: Timber.
Classen-Bockhoff R., Froebe H. A. & Langerbeins D. 1989: Die Infloreszenzstruktur von Gundelia tournefortii L. (Asteraceae). – Flora 182: 463-479.
Jeffrey C. 1966: Notes in Compositae I. The Cichorieae in East Tropical Africa. – Kew Bulletin 18: 427-486.
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.
Lack H. W. 2006: Tribe Cichorieae Lam. & DC. – Pp. 180-199 in: Kadereit J. W. & Jeffrey C. (ed.), The families and genera of vascular plants 8. – Berlin: Springer.
Stebbins G. L. 1953: A new classification of the tribe Cichorieae, family Compositae. – Madroño 12: 65–81.
Tremetsberger K., Gemeinholzer B., Zetzsche H., Blackmore S., Kilian N. & Talavera S. 2012: Divergence time estimation in Cichorieae (Asteraceae) using a fossil-calibrated relaxed molecular clock. – Org. Divers. Evol. 13: 1-13. // ➪ //