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Alla Alla använda. att fritt och är universitetsbibliotek Göteborgs vid digitaliserats har verket Det här söka och kan kopiera du Det text. betyder att till maskinläsbar OCR-tolkade tryckta är texter Johan Rova Botanical Institute Göteborg University Sweden 1999 Gh9 The Condamineeae-Rondeletieae-Sipaneeae Complex (Rubiaceae) llONDELETIEAE SIPANEEAE COMPLEX iölå) % S I Biblioteket för botanik och miljö Göteborg University Faculty of Natural Sciences Botanical Institute Dissertation THE RONDELETIEAE-CONDASVUNEEAE-SIPANEEAE COMPLEX (RUBIACEAE) Johan H. E. Rova jfpsyp IIS«! Göteborg 1999 i? ' K"-'-* Ä0||KS>"vS Ö>xl J • ' f 1.4 . 8H ... ÏW• Ää? ;; liHi fÉÉI . ::t# • • • 1 sk - • I. s.i: •'>:' 'Vs . ' V ff •• '-vvi.•' :i ' fe; Ä ' • ' I 'î<\ 1 -Ii .:»,. vpr' ' r&j : a V: SI' ' :: §||S#S H . — , g,"# •Î , -twW/\ s B•i m Iii!!SU|§|1| •^m;|MS;:ÊS»4 «2] 1;'v,,. •• ••, . , ;...c:', ... S TI'' • '. ' ..-*• ' ' "Int IIIilÄfSll : • :•••.'•'• S* S i,•?^'; v'4--'•>-r--* jail h- - s• '. äfvÄÄ sa Ü , : • B S " v'vl'lvV ' ,%• j 4 : ' • 1 ' '• m ,. i-v .4''-. 1} w • ;•' " . •' .-.-v.v - . 4- fk St ' j : : ; • vi-S-v '.'1;l". .''l''.l- "öiv '?si .' ' V 'V S " V;K t' ''-v. Errata J.H.E. Rova. 1999. The Condamineeae-Rondeletieae-Sipaneeae Complex (Rubiaceae). Botanical Institute, Göteborg University, Box 461, SE-405 30 Göteborg. 1. The title on the cover pages should read "The Condamineeae-Rondeletieae-Sipaneeae Complex (Rubiaceae)". 2. Appendix 2 in Paper II was left out by the printing office and should read: APPENDIX 2: Classification history Subfamilial and tribal classifications of analyzed Rubiaceae genera as previously proposed by various authors, and compared to the results of the present rps\6 phylogeny. Only genera not listed in Rova et al. (submitted) are included. Subfamilies ("tribes" in Candolle, 1830) are given with four letters: anti-Antirheoideae, cinc- Cinchonoideae, coff-Coffeoideae, gard-Gardeniaceae, guet-Guettardoideae, hame-Hamelieae, hame- Hamelieae, hedy-Hedyotideae, hill-Hillioideae, ixor-Ixoroideae, oper-Opercularieae, and rubi-Rubioideae. Tribes ("subtribes" in Candolle, 1830) are given with three letters: chi-Chiococceae, cin-Cinchoneae, con- Condamineeae, cop-Coptosapeltae, gar-Gardenieae, gue-Guettardeae, ham-Hamelieae, hil-Hillieae, nau- Naucleeae, and ron-Rondeletieae. Hooker's "series" are given with one letter: a-Series A (many ovules in each locule) and c-Series C (solitary ovule in each locule). means that the genus (or a synonym for it) was not considered by the author; "?" means an uncertain position according to the author. Footnotes in the table are as follows:a as Laugeria,b as Anthocephalus,c implicitly in that Bremekamp is supposed to follow 1) Schumann and 2) Hooker in this case,d as included in Calycophyllum. Genus Candolle Hooker Schu­ Verdcourt Breme­ Robbrecht Robbrecht rps16 (1830) (1873) mann (1958) kamp (1988) (1993) (1891) (1966) Alibertia hame a-gar cinc-gar ixor-gar ixor-gar ixor-gar IX02 Amaioua gard-gar a-gar cinc-gar - ixor-gar ixor-gar IX02 Atractogyne cinc-gar ixor-gar ixor-gar IX02 Bobea Gaud. c-gue coff-gue guet-guec anti-gue anti-gue CIN5 Bothriospora a-ham cinc-gar ? ? IX01 Burchellia gard-gar a-gar cinc-gar ixor-gar ixor-gar IX02 Cremaspora c-alb coff-alb cinc-ixo - ixor-gar ixor-gar IX02 Cuatrecasasiodendron cinc-ron cinc-ron CIN5a Deppea oper a-ron eine-ron rubi-? ? rubi-ham rubi-ham CIN2 Dolicholobium a-cin cinc-cin cinc-cin cinc-cin IX01 ixor-oct Fernelia gard-gar a-gar cinc-gar ixor-oct IX02 (as hyp) Hillia cinc-cin a-cin cinc-cin rubi-? hill-hil cinc-hil cinc-hil CIN2 not Hymenodictyon cinc-cin a-cin cinc-cin cinc-cin cinc-cin cinc-cop CIN3 ixor-cop Kerianthera cinc-con cinc-con CIN6 Macrocnemum P. Br. hedy-ron a-cin cinc-cin cinc-cin cinc-cin IX01 Malanea guet-gue c-gue coff-gue • anti-gue anti-gue CIN5a Mastixiodendron ?anti-chi anti-?chi IX01 Neolamarckia cinc-?cinb cinc-nau cinc-nau CIN3 Neolaugeria c-gue* anti-gue anti-gue CIN5a Oxyanthus gard-gar a-gar cinc-gar ixor-gar ixor-gar IX02 Sabicea hame-ham a-ise cinc-mus cinc-mus cinc-sab cinc-ise cinc-ise IX05 Semaphyllanthe cinc-cind cinc-cin" IXOi Stilpnophyllum a-cin cinc-cin cinc-cin cinc-?cin CIN1 Virectaria hedy-ron a-hed - cinc-7 urop-oph rubi-hed rubi-hed IX05 30 - - . Göteborg University Faculty of Natural Sciences Botanical Institute Dissertation THE RONDELETIEAE-CONDAMINEEAE-SIPANEEAE COMPLEX (RUBIACEAE) Johan H. E. Rova Göteborg 1999 Göteborg University Faculty of Natural Sciences Botanical Institute Dissertation for the Degree of Doctor of Philosophy in Systematic Botany presented at Göteborg University, Botanical Institute, Carl Skottsbergs gata 22b, Göteborg, at 10.00, 26 November, 1999. ABSTRACT Rova, J. H. E. 1999. The Condamineeae-Rondeletieae-Sipaneeae complex (Rubiaceae). Botanical Institute, University of Göteborg, Sweden. ISBN 91-88896-16-1. Tribal demarcations in the Condamineeae-Rondeletieae-Sipaneeaecomplex (Rubiaceae) are inferred from phylogenies based on chloroplast trnL intron, trnL-F spacer, and rps16 DNA sequence data. Cladistic analyses of the trnL-F data and rps 16 data are presented in two separate papers, and a jackknife analysis of the combined data set (including126 Rubiaceae terminals and 738 informative characters) is presented in the thesis. It is found that the Condamineeae-Rondeletieae-Sipaneeae complex is not monophyletic. Condamineeae in a strict sense forms a poorly resolved complex together with Calycophylleae, Hippotideae, and Simireae in subfamily Ixoroideae. Rondeletieae in a strict sense belongs in subfamily Cinchonoideae. Sipaneeae is found in subfamily Ixoroideae. Anumber of genera should be transferred from Rondeletieae to the Condamineeae complex and other parts of subfamily Ixoroideae. Other genera should be transferred from Rondeletieae to Guettardeae, which is found to be the sister tribe to Rondeletieae. Support was found for a splitting of Rondeletia into several smaller genera, but the generic boundaries between some of these segregates could not be finally settled. The Portlandia-group is closely related to Catesbaeeae and Chiococceae within Cinchonoideae. Limited support isfound for aseparation of Catesbaeeae from Chiococceae.Further, subfamily Antirheoideae is shown to be polyphyletic, and only three large subfamilies should be recognized in Rubiaceae: Cinchonoideae, Ixoroideae, and Rubioideae. The taxonomic history of the tribes of central importance to this study is briefly reviewed, and systematic positions are suggested for a number of hitherto unplaced genera. KEY WORDS Antirheoideae, Catesbaeeae, Chiococceae, Condamineeae, Guettardeae, Rondeletieae, rps 16, Rubiaceae, Sipaneeae, trnL-F Johan Rova, Botanical Institute, Göteborg University, Box 461, SE-405 30 Göteborg, Sweden Botanical Institute, Göteborg ISBN 91-88896-16-1 Printed by Vasastadens Bokbinderi AB, Västra Frölunda, Sweden, 1999. ' ' The Condamineeae-Rondeletieae-Sipaneeae Complex (Rubiaceae) Johan H. E. Rova INTRODUCTION grown for ornamental purposes (e.g., Gardenia and Ixora), or as dyes( Rubia) and tanning agents To communicate, we need a common language. (Uncaria) (Aldén et al., 1998). A number of In this language, we must agree on what to call Rubiaceae species are Ni-hyperaccumulators the different objects and phenomena that we find attaining a dry matter Ni concentration of 1-5% interesting enough to discuss with other people. (Jaffré and Schmid, 1974; Reeves et al., 1999). To understand the world around us, we must also establish how these objects and phenomena are A family the size of the Rubiaceae needs to be related to each other - both physically and subdivided into smaller and more manageable causally. In biology, as well as in other sciences, units in order to facilitate the understanding of the naming of things is the field of taxonomy, its diversity and evolution. Robbrecht (1988) whereas the establishment of relationships is the elegantly presented a modern view of Rubiaceae field of systematics. Taxonomy and systematics interrelationships by drawing tribes and are interconnected. In modern biology, a sound subfamilies as circles and borders in a two- taxonomy should mirror the evolutionary dimensional landscape. The tribes relationships of a group, i.e., we should endeavor Condamineeae, Rondeletieae, and Sipaneeae to classify groups of organisms (taxa) based on were drawn as confluent circles within subfamily their common ancestry, not on how similar they Cinchonoideae. By doing this, he stressed that look. Such a sound taxonomy is called natural. tribal demarcations in this complex were in need A natural taxonomy is the necessary basis for all of a more thorough investigation. studies where interest is put on more than one This thesis work started out as an attempt to individual—be that character evolution, ecology, find tribal demarcations within this conservation biology, or the simple wish to find predominantly neotropical Condamineeae- relatives to a medically interesting plant. The aim Rondeletieae-Sipaneeae complex. As the study of this study is to provide a phylogeny for the progressed, however, unexpected results establishment of a natural and practically useful appeared and a number of other tribes also had taxonomy within the plant family Rubiaceae.