Selbyana 31(2): 68–94. 2013. ANEW FORMAL CLASSIFICATION OF GESNERIACEAE ANTON WEBER* Department of Structural and Functional Botany, Faculty of Biodiversity, University of Vienna, A-1030 Vienna, Austria. Email: [email protected] JOHN L. CLARK Department of Biological Sciences, The University of Alabama, Tuscaloosa, AL 35487, USA. MICHAEL MO¨ LLER Royal Botanic Garden Edinburgh, Edinburgh EH3 5LR, Scotland, U.K. ABSTRACT. A new formal classification of Gesneriaceae is proposed. It is the first detailed and overall classification of the family that is essentially based on molecular phylogenetic studies. Three subfamilies are recognized: Sanangoideae (monospecific with Sanango racemosum), Gesnerioideae and Didymocarpoideae. As to recent molecular data, Sanango/Sanangoideae (New World) is sister to Gesnerioideae + Didymocarpoideae. Its inclusion in the Gesneriaceae amends the traditional concept of the family and makes the family distinctly older. Subfam. Gesnerioideae (New World, if not stated otherwise with the tribes) is subdivided into five tribes: Titanotricheae (monospecific, East Asia), Napeantheae (monogeneric), Beslerieae (with two subtribes: Besleriinae and Anetanthinae), Coronanthereae (with three subtribes: Coronantherinae, Mitrariinae and Negriinae; southern hemisphere), and Gesnerieae [with five subtribes: Gesneriinae, Gloxiniinae, Columneinae (5the traditional Episcieae), Sphaerorrhizinae (5the traditional Sphaerorhizeae, monogeneric), and Ligeriinae (5the traditional Sinningieae)]. In the Didymocarpoideae (almost exclusively Old World, especially E and SE Asia/Malesia) two tribes are recognized: Epithemateae [with four small, but morphologically and genetically very distinctive subtribes: Loxotidinae (monogeneric with Rhynchoglossum), Monophyllaeinae, Loxoniinae and Epithematinae (monogeneric)] and Trichosporeae (the earliest name at tribal rank for the ‘‘Didymocarpoid Gesneriaceae’’). The last is subdivided into ten subtribes: Jerdoniinae (monospecific), Corallodiscinae (monogeneric), Tetraphyllinae (monogeneric), Leptoboeinae, Ramondinae (Europe), Litostigminae (monogeneric), Streptocarpinae (Africa and Madagascar), Didissandrinae, Loxocarpinae and Didymocarpinae. Didymocarpinae is the largest subtribe (ca. 30 genera and .1600 species) and still requires intensive study. It includes the most speciose genera such as Cyrtandra, Aeschynanthus, Agalmyla, Didymocarpus, Henckelia, Codonoboea, Oreocharis and Primulina and the types of the traditional tribes Didymocarpeae, Trichosporeae and Cyrtandreae. Key words: Gesneriaceae, classification, traditional classifications, molecular systematics INTRODUCTION Since Weber’s (2004a) treatment, many more molecular studies have been published (reviewed in The incorporation of molecular methods into Mo¨ller & Clark, 2013). In the New World plant systematics over the last two decades has Gesneriaceae a new tribe has been erected (Roalson dramatically changed our understanding of the et al. 2005b), and in the Old World Gesneriaceae a phylogenetic diversification of angiosperms. Not much more detailed picture of the informal groups unexpectedly, this applies also to the family previously recognized emerged through the work of Gesneriaceae. The last classification, based on M. Mo¨ller and his collaborators (e.g., Mo¨ller et al. morphological (and with respect to the Neotropical 2009, 2011a; Weber et al. 2011a). This paper is the Gesneriaceae also cytological) characters was that of first to propose a comprehensive classification of the Burtt and Wiehler (1995). This was followed by the entire family based on molecular phylogenetic treatment of the family in Kubitzki’s ‘‘Families and studies and to formalize a rank-based system from genera of vascular plants’’ by Weber (2004a). In the published phylogenies. latter work, allowance was made for the molecular data then available (partly unpublished and pub- lished later by Mo¨ller et al. 2009). This resulted in an DO WE NEED FORMAL RANK-BASED abandonment of the traditional tribes hitherto CLASSIFICATIONS? recognized in the Old World Gesneriaceae. For a provisional subdivision of the ‘‘Didymocarpoid The answer is: not necessarily. Formal classifi- Gesneriaceae,’’ informal group names (e.g., ‘‘Basal cations with ranks indicated by name endings are Asiatic genera’’) were used instead of formal names. simply a well-established custom. The modern language of our discipline consists of precise and * Corresponding author. easily comprehensible communication via phylograms 68 Selbyana selb-31-02-02.3d 10/12/13 04:38:30 68 WEBER ET AL.: NEW CLASSIFICATION OF GESNERIACEAE 69 or cladograms (‘‘phylogenetic trees’’). Trees are Lamiales. The inflorescences are said to be graphical representations (branching diagrams) of racemose, and thus are abnormal in the Old the relationships of taxa and can do without names World Gesneriaceae. Burtt (1963: 217) wrote that that reflect the hierarchical order. Where complex the genus ‘‘is at present best regarded as a slightly diversification patterns exist, they provide more anomalous member of Scrophulariaceae, … whose precise information than formal classifications limits need a thorough overhaul.’’ In the meantime, (with a limited number of ranks) could ever do. Scrophulariaceae have been split into many Traditional rank-based classifications date back families of their own (e.g., Olmstead 2001, to a time when nobody could imagine how precise Albach et al. 2005, Oxelman et al. 2005, Tank et phylogenetic reconstructions would ever be possi- al. 2006, APG III 2009, Scha¨ferhoff et al. 2010). ble. Therefore, only few (three) formal ranks Fischer (2004) referred Brookea to the between family and genus have been established: Scrophulariaceae-‘‘Stilbaceae.’’ To the best of our subfamily, tribe and subtribe. Reveal (1995 onw, knowledge there is no molecular data available for edition 2011; 2010) has suggested three additional the genus (see also Wolfe et al. 2006). Brookea is ranks be implemented below the family level clearly in need of a detailed morphological and (supersubfamily, supertribe, supersubtribe). In ear- molecular study. At present, there is no specific lier versions of this manuscript we attempted to evidence that it does belong to Gesneriaceae. introduce some of these new ranks. However, due to conflicting or incomplete molecular evidence, Charadrophila Marloth. The systematic position the relationships of the subtribes within the large of this South African monotypic genus has been tribes Gesnerieae and Trichosporeae (in the new disputed since its establishment by Marloth (1899), sense, see below) proved ambiguous. Therefore, we having been moved between Scrophulariaceae and refrain from establishing a complex hierarchical Gesneriaceae. Based on a detailed morphological classification beyond the traditional ranks. and ontogenetical study, Weber (1989) reached the Systematics is a ‘‘never-ending synthesis’’ and conclusion that Charadrophila should be removed time will perhaps never be ripe for an ultimate from Gesneriaceae and placed in the Scrophulariaceae classification. However, at present we have (s.l.). Fischer (2004) placed the genus in Scrophu- molecular data of all major subgroups of Gesner- lariaceae (s.str.) tribe Alonsoeae. More recently, and iaceae and an updated classification along the lines based on molecular data, Charadrophila was placed addressed here appears meaningful. The classifi- in Stilbaceae, a segregate family of traditional cation of the neotropical Gesneriaceae seems to Scrophulariaceae comprising mainly ‘‘scrophulari- approach consolidation, and the recent phyloge- aceous’’ genera from the Western Cape (Oxelman et netic analyses of the Old World Gesneriaceae al. 2005, APG III 2009). without a doubt are a major step forward (see review of Mo¨ller & Clark 2013). Thus, the Cubitanthus Barringer. From collections foundation is laid to synthesize a new and overall examined by the authors and photographs taken classification of the family. by A. Chautems (Gene´ve), Cubitanthus [mono- typic with C. alatus (Cham. & Schltdl.) Barringer, WHAT TO DO WITH ‘‘ANOMALOUS’’ GENERA? originally described as Russelia alata by Chamisso and Schlechtendal (1828: 3), and referred to Scro- Inclusion of a number of genera in the phulariaceae] is clearly a member of Scrophu- Gesneriaceae has been debated (see Weber lariaceae (s.l.) and must be excluded from 2004a, under ‘‘Genera of uncertain familial Gesneriaceae where the species was placed by affiliation’’ and ‘‘Excluded genera’’). All may Bentham (1876, as Russelia alata) and Barringer be, and some definitely are, of relevance for the (1984, as Cubitanthus alatus). Recent molecular classification of the family. In the following, the data (Perret et al. 2013) support the placement of genera are listed (in alphabetical sequence) along Cubitanthus in or close to Linderniaceae and not in with brief comments. The molecular aspects have Gesneriaceae. been addressed in more detail by Mo¨ller and Clark (2013). Cyrtandromoea Zoll. The genus was established by Zollinger (1854–1855), with the name Brookea Benth. This is a genus of four species reminiscent of Cyrtandra J.R.Forst. & G.Forst., of shrubs or small trees from Borneo. It was placed the largest genus of the Didymocarpoid Gesne- in Scrophulariaceae by Bentham (1876), but riaceae. In early classifications (Bentham 1876, referred to Gesneriaceae by Hallier (1903). The Clarke 1883, Fritsch 1893–1894, Ridley 1923), flower has some similarity with a gesneriad flower, Cyrtandromoea was always