TRIBES AND GENERA OF THE Lennart Andersson2CINCHONEAE COMPLEX(RUBIACEAE)'ABSTRACT The phylogenetic relationships of taxa recently referred to the tribes Calycophylleae, Cinchoneae, and Hillieaewere analyzed using cladistic methods. The analyses used morphological and micromorphological characters, and aselection of 60 species were used as terminal taxa. It was concluded that ail three tribes are supported as monophyleticbut must be altered somewhat in circumscription. The tribes Cinchoneae and Hillieae are sister groups, and thesetwo in turn form the sister group of the Calycophylleae. The tribe Calycophylleae should comprise the genera Alseis,Calycophyllum, Capirona, Ferdinandusa, Macrocnemum, Wittmackanthus, and the new genus SemaphyllantheL. Andersson here segregated from Calycophyllum. These genera are ail supported as monophyletic, although thesupport is sometimes weak. The tribe Hillieae should comprise Balmea, Blepharidium, Cosmibuena, and Hillia. Itwas not tested whether or not these genera are monophyletic. The tribe Cinchoneae should comprise Cinchona,Joosia, Ladenbergia, Maguireocharis, Pimentelia, Remijia, Stilpnophyllum, and the new genus Cinchonopsis L.Andersson here erected for Cinchona amazonica. Dolicholobium occurs as a basal clade of the Cinchoneae but itsposition is unstable and it is possibly not related to this alliance at ail. Cinchona is supported as monophyletic if C.amazonica and C. henleana are excluded, the latter one being transferred to Ladenbergia. Ladenbergia is supportedas monophyletic in approximately the traditional sense. Remijia is supported as monophyletic if Cephalodendron isincluded. Joosia and Stilpnophyllum are supported as monophyletic, while the remaining genera are monotypic. Sixnew combinations are made: Semaphyllanthe intonsa (Steyerm.) L. Andersson, S. megistocaula (K. Krause) L.Andersson, S. merumensis (Steyerm.) L. Andersson, S. obovata (Ducke) L. Andersson, S. spectabilis (Steyerm.) L.Andersson, S. venezuelensis (Steyerm.) L. Andersson, and Cinchonopsis amazonica (Standley) L. Andersson. The tribe Cinchoneae is often widely circum-scribed (see, e.g., de Candolle, 1830; Schumann,1891, 1897; Robbrecht, 1988). In this sense it isa pantropical group comprising ail genera withmultiovulate ovaries, ascendingly imbricate ovules,and winged seeds. The case was examined by An-dersson & Persson (1991), who concluded that anumber of genera traditionally included in the Cin-choneae are not at ail closely related to Cinchona.Out of 50 genera referred to the Cinchoneae byRobbrecht (1988), 10 were transferred to the Hed-yotideae and six to the Condamineae. Disregardingfour genera left as incertae sedis, the remainderwere divided among four tribes. The paleotropicalgenera, except Dolicholobium, were referred tothe tribe Coptosapelteae. Four genera, Alseis, Ca-lycophyllum, Schizocalyx, and Wittmackanthus,were referred to the new tribe Calycophylleae. Hil-lia was left in a monotypic tribe of its own, theHillieae. Thirteen genera were left in the Cincho-neae: Capirona, Cephalodendron, Cinchona,Cosmibuena, Dolicholobium, Ferdinandusa,Joosia, Ladenbergia, Macrocnemum, Maguireo-charis, Pimentelia, Remijia, and Stilpnophyl-' This study was supported by a grant (B-BU 08997Research Council. 2 Dept. of Systematic Botany, Univ. of GCteborg, Carl lum. The relationships between these four tribes were not resolved in strict consensus although a majority of equally parsimonious solutions sug-gested that the Coptosapelteae are possibly the sister group of the three neotropical tribes, which in turn may form a monophyletic group. The tribes Cinchoneae and Hillieae are probably sister groups, forming together the sister group of the Calyco-phylleae. The investigation of Andersson and Persson was based in large part on data from the literature, using genera as terminal taxa. The aim of the present study is to reach higher resolution regard-ing circumscription and interrelationships of the neotropical tribes and genera of the Cinchoneae complex, using original observations throughout and species as terminal taxa. Two initial assumptions are based on the findings of Andersson & Persson: (1) the neotropical tribes Calycophylleae, Cinchoneae, and Hillieae form a monophyletic group, possibly including also the genera Balmea and Blepharidium; (2) the sister group of this complex is the tribe Coptosapelteae. Neither of these assumptions was fully supported,-307 to L. Andersson) from the Swedish Natural ScienceSkottsbergs Gata 22, S-413 19 Giteborg, Sweden.N. MISSOURI BOT. GARD. 82: 409-427. 1995.
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