1547 Alejandro & al. • Classification and revision of Greeniopsis TAXON 59 (5) • October 2010: 1547–1564 INTRODUCTION Greeniopsis Merr., established by Merrill (1909), is a poorly known genus of Rubiaceae endemic to the Philippines. The genus at present comprises seven species ranging from shrubs to medium-sized trees: G. discolor Merr., G. euphle- bia Merr., G. megalantha Merr., G. multiflora (Elmer) Merr., G. philippinensis Merr., G. pubescens Merr., and G. sibuya- nensis Elmer. Greeniopsis is characterized by terminal in- florescences, contorted corolla aestivation, bilocular ovaries with elongated placentas, capsular fruits, winged seeds, and 3-colporate pollen (Tange, 1996b). Although the genus without doubt belongs to Rubiaceae (opposite-decussate leaves, presence of stipules, actinomorphic flowers, inferior ovary), its tribal position within this family remains unsettled. Merrill (1909) proposed a close relation of Greeniopsis to the Asian genera Emmenopterys Oliv. (two spe- cies) and Mussaendopsis Baill. (four species). A relationship between Greeniopsis and Mussaendopsis was also put forward by Krause (1915) based on their contorted aestivation, bilocular ovaries, capsular fruits, winged seeds and elongated placentas. Later, Robbrecht (1988) placed Greeniopsis as well as Em- menopterys and Mussaendopsis in Cinchoneae-Cinchoninae, based on Krause’s work (E. Robbrecht, pers. comm.). Subse- quent morphological studies tentatively transferred Greeni- opsis and Mussaendopsis to Coptosapelteae (Andersson & Persson, 1991; Robbrecht, 1993). Recent molecular data later demonstrated that both Emmenopterys and Mussaendopsis belong to the Calycophylleae/Condamineeae alliance of Ix- oridinae I (Robbrecht & Manen, 2006; Bremer & Eriksson, 2009). However, because no molecular data were available for Greeniopsis, its taxonomic position remained unresolved after the dismantling of the Coptosapelteae and the transfer of most of its genera to Naucleeae in the wide sense (Razafimandim- bison & Bremer, 2002). In contrast to Merrill (1909) and later authors, Tange (1996a,b) disagreed on the close affinity between Greeniopsis and Mussaendopsis mainly because of their different stipule form, inflorescence position, and floral morphology (Puff & Igersheim, 1994). Moreover, Mussaendopsis is reminiscent, hence its name, of Mussaenda Burm. ex L. in possessing en- larged calyx lobes or semaphylls (sensu Leppik, 1977), which are absent in Greeniopsis. Instead, Tange (1996a,b) postulated a relationship of Greeniopsis with two other Southeast Asian genera, Aleisanthia Ridl. (two species) and Aleisanthiopsis Tange (two species). The three genera share a similar contorted corolla, secondary pollen presentation, idioblasts with a single druse, elongated placentas, and capsular fruits with hard en- docarp. Following Ridley (1920), Tange (1996b) suggested the placement of Aleisanthia and Aleisanthiopsis in Rondeletieae but refrained commenting on the tribal position of Greeniopsis. Plastid data ( rps16, rbcL, trnL-F ) have disclosed a re- lationship between Aleisanthia-Aleisanthiopsis, the South- east Asian genus Greenea Wight & Arn. (ca. 9 species) and the large pantropical genus Ixora L. (ca. 400 species) (Rova & al., 2002; Mouly & al., 2009a). Accordingly, Mouly & al. Molecular support of the classification of Greeniopsis Merr. in Aleisanthieae (Rubiaceae), with a revision of the genus Grecebio Jonathan D. Alejandro, 1,2,3 Ulrich Meve, 3 Millard Uy, 1 Arnaud Mouly, 4 Mike Thiv 5 & Sigrid Liede-Schumann 3 1 College of Science and 2 Research Center for the Natural Sciences, University of Santo Tomas, España Boulevard, 1015 Manila, Philippines 3 Department of Plant Systematics, University of Bayreuth, Universitätstr. 30, 95440 Bayreuth, Germany 4 Université de Franche-Comté – CNRS UMR 6249 Chrono-environnement, UFR Sciences et Techniques, 16 Route de Gray, 25030 Besançon cedex, France 5 Botany Department, Museum of Natural History Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany Author for correspondence: Grecebio J.D. Alejandro, gdalejandro@mnl.ust.edu.ph Abstract Greeniopsis Merr. is a small genus of Rubiaceae endemic to the Philippines. Previously, no molecular data have been available for Greeniopsis and its taxonomic position has remained controversial. Based on morphology, the genus was tentatively included in the tribe Aleisanthieae (Ixoroideae) along with the Southeast Asian genera Aleisanthia and Aleisanthiopsis. To determine the taxonomic position of Greeniopsis and evaluate the monophyly of the genus, sequences of the rps16 and trnT-F regions of cpDNA of all seven Greeniopsis species were generated. Parsimony and Bayesian analyses of the combined plastid dataset strongly support the inclusion of Greeniopsis in Aleisanthieae as well as its monophyly. Woolly hairs on the abaxial leaf surface occur in only one Greeniopsis species and can no longer be regarded a synapomorphy of the tribe. The generic status of Aleisanthia, Aleisanthiopsis, and Greeniopsis is maintained. A revision of Greeniopsis, including a key to species, descriptions, distribution maps, and illustrations, is provided. Keywords Aleisanthia ; Aleisanthieae; Aleisanthiopsis; Greeniopsis; Ixoroideae; Philippines; rps16; Rubiaceae; trnT-F