801 Mouly & al. • Phylogeny of the Gardenieae complex (Rubiaceae) TAXON 63 (4) • August 2014: 801–818 Version of Record (identical to print version). Received: 28 Dec 2012 | returned for first revision: 10 Jul 2013 | last revision received: 27 May 2014 | accepted: 27 May 2014 | not published online ahead of inclusion in print and online issues || © International Association for Plant Taxonomy (IAPT) 2014 INTRODUCTION Rubiaceae is the fourth-largest flowering plant fam- ily (Davis & al., 2009), although only a few species are well known and commonly used, such as coffee and the fragrant ornamental ixoras and gardenias. Coffea L., Ixora L., and Gardenia J.Ellis (respectively) belong to the same Rubiaceae lineage, Ixoroideae, a subfamily with a long and intricate tax- onomic history and an unsatisfactorily resolved phylogeny, despite numerous recent studies (e.g., Persson, 1996, 2000a; Andreasen & Bremer, 2000; Rova & al., 2002; Robbrecht & Manen, 2006; Bremer & Eriksson, 2009; Razafimandim- bison & al., 2011; Kainulainen & al., 2013). Within Ixoroideae, tribe Gardenieae is a large and morphologically diverse group of ca. 100 genera (Fig. 1), whose circumscription has long been controversial (e.g., Candolle, 1830; Schumann, 1891; Verdcourt, 1958; Bremekamp, 1966; Robbrecht & Puff, 1986; Andreasen & Bremer, 1996, 2000; Robbrecht & Manen, 2006; see Table 1). Candolle (1830) described Gardenieae to accommodate Rubi- aceae genera with indehiscent fleshy bilocular (rarely uniloc- ular) fruits and “remarkable” flowers (the author meant large flowers) with or without pedicel. The original circumscription (Candolle, 1830) included 28 genera: Sarcocephalus Afzel. ex R.Br., Zuccarinia Blume, Schradera Vahl (as Lucinaea DC.), Burchellia R.Br., Amaioua Aubl., Mussaenda L. (incl. Mene- storia DC.), Kutchubea Fisch. ex DC., Isertia Schreb. (as Cas- supa DC.), Aidia Lour. (as Gynopachis Blume, Cupia DC., and Stylocoryna Cav.), Tocoyena Aubl., Posoqueria Aubl., Oxyan- thus DC., Genipa L., Gardenia, Randia L. (incl. Oxyceros and Euclinia ), Chapelieria A.Rich., Heinsia DC., Timonius DC. (as Phylogenetic structure and clade circumscriptions in the Gardenieae complex (Rubiaceae) Arnaud Mouly, 1 , 2 Kent Kainulainen, 1 Claes Persson, 3 Aaron P. Davis, 4 Khoon Meng Wong, 5 Sylvain G. Razafimandimbison1 & Birgitta Bremer1 1 The Bergius Foundation, The Royal Swedish Academy of Sciences, and Department of Ecology, Environment and Plant Sciences, Stockholm University, 106 91, Stockholm, Sweden 2 Université de Franche-Comté—CNRS UMR 6249 Chrono-environnement, ESPE de Franche-Comté, 16 Route de Gray, 25030 Besançon cedex, France 3 Department of Biological and Environmental Sciences, University of Gothenburg, P.O. Box 461, 405 30, Göteborg, Sweden 4 Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, U.K. 5 Singapore Botanic Gardens, 1 Cluny Road, Singapore 259569 Author for correspondence: Arnaud Mouly, arnaud.mouly@univ-fcomte.fr DOI http://dx.doi.org/10.12705/634.4 Abstract In this study we investigate the large and diverse Rubiaceae-Gardenieae and closely related tribes Bertiereae, Coffeeae, Cremasporeae, Octotropideae, and Pavetteae. Some of the tribes or groups have been shown to be monophyletic and strongly supported, but the phylogeny of this large complex is still far from being satisfactorily elucidated particularly for Gardenieae, both in terms of intertribal relationships as well as tribal delimitations. We reconstruct the phylogeny of the complex using an extensive sampling of 108 genera and five plastid DNA regions. Phylogenetic relationships demonstrate that Gardenieae sensu Andreasen & Bremer is polyphyletic, as Burchellia, Didymosalpinx, Monosalpinx, and Mantalania are closer to Octotropideae- Cremasporeae. In addition, Pavetteae and the investigated members of Aulacocalyceae are nested in a supported but partially unresolved Gardenieae-Pavetteae clade. Within this clade, several strongly supported groups are resolved: an Aidia group, an Alibertia group, a Gardenia group, Pavetteae including Pelagodendron, a Porterandia group, a Randia group, a Rothmannia group (including Aulacocalyx and Heinsenia), a Sherbournia group, and the two isolated genera Massularia and Schuman- niophyton. The latter genus presented a high rate of genetic substitutions, which resulted in perturbations of the phylogenetic reconstruction. A revised tribal circumscription is given for Gardenieae, the Alibertia group is recognized at tribal level as an emended Cordiereae, and a new tribe, Sherbournieae, is described to accommodate the members of the Sherbournia group. Keywords Aulacocalyceae; Cordiereae; Gardenieae; mapping; molecular phylogeny; Octotropideae; Pavetteae; radiation; Rubiaceae; Sherbournieae Supplementary Material Electronic Supplement (Fig. S1) and alignment are available in the Supplementary Data section of the online version of this article at http://www.ingentaconnect.com/content/iapt/tax