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Rubiaceae) Necessitated by the Polyphyly of Bikkia Reinw

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Rubiaceae) Necessitated by the Polyphyly of Bikkia Reinw Reinstatement of the endemic New Caledonian genus Thiollierea Montrouz. (Rubiaceae) necessitated by the polyphyly of Bikkia Reinw. as currently circumscribed Laure BARRABÉ Institut de Recherche pour le Développement, UMR AMAP, Laboratoire de Botanique et d’Écologie végétale appliquées, Herbarium NOU, BPA5, F-98848 Nouméa (New Caledonia) [email protected] Arnaud MOULY Université de Franche-Comté, UMR CNRS 6249 Chrono-Environnement, IUFM de Franche-Comté, 16 route de Gray, F-25030 Besançon (France) [email protected] Porter P. LOWRY II Missouri Botanical Garden, P.O. Box 299, St. Louis, MO, 63166-0299 (USA) [email protected] and Muséum national d’Histoire naturelle, Département Systématique et Évolution, UMR 7205, case postale 39, 57 rue Cuvier, F-75231 Paris cedex 05 (France) [email protected] Jérôme MUNZINGER Institut de Recherche pour le Développement, UMR AMAP, Laboratoire de Botanique et d’Écologie végétale appliquées, Herbarium NOU, BPA5, F-98848 Nouméa (New Caledonia) [email protected] Barrabé L., Mouly A., Lowry II P. P. & Munzinger J. 2011. — Reinstatement of the endemic New Caledonian genus Thiollierea Montrouz. (Rubiaceae) necessitated by the polyphyly of Bikkia Reinw. as currently circumscribed. Adansonia, sér. 3, 33 (1) : 115-134. DOI: 10.5252/ a2011n1a8. ABSTRACT Th e genus Bikkia Reinw. as currently circumscribed comprises 20 species dis- tributed throughout the western Pacifi c Ocean, with a center of diversity in New Caledonia (11 species, all but one endemic). Two recent phylogenetic studies based respectively on molecular and morphological data have shown that Bikkia s.l. is polyphyletic, comprising two lineages: coastal Bikkia from the western Pacifi c (including the type species of the genus, B. tetrandra (L.f.) A.Rich.) and endemic New Caledonian Bikkia, a situation that necessitates the resurrection ADANSONIA, sér. 3 • 2011 • 33 (1) © Publications Scientifi ques du Muséum national d’Histoire naturelle, Paris. www.adansonia.com 115 Barrabé L. et al. of the generic name Th iollierea Montrouz., the oldest available for the second KEY WORDS Rubiaceae, of these groups. An emended description is provided for Th iollierea as well as Bikkia, a summary of the features that distinguish it from Bikkia s.s., and seven new Th iollierea, New Caledonia, combinations are proposed (T. kaalaensis (N.Hallé & Jérémie) Barrabé & Mouly, endemic genus, T. lenormandii (N.Hallé & Jérémie) Barrabé & Mouly, T. neriifolia (Brongn.) polyphyly, Barrabé & Mouly, T. pachyphylla (Guillaumin) Barrabé & Mouly, T. parvifl ora taxonomic reinstatement, (Schltr. & K.Krause) Barrabé & Mouly, T. retusifl ora (Brongn.) Barrabé & new combinations. Mouly and T. tubifl ora (Brongn.) Barrabé & Mouly). RÉSUMÉ Rétablissement taxonomique du genre Th iollierea Montrouz. (Rubiaceae) endé- mique de Nouvelle-Calédonie, nécessité par la polyphylie de Bikkia Reinw. tel qu’actuellement défi ni. Le genre Bikkia Reinw. s.l. tel qu’il est défi ni actuellement comporte 20 espèces réparties sur l’ensemble du Pacifi que ouest. Son centre de diversité se trouve en Nouvelle-Calédonie avec 11 espèces, dont 10 endémiques de l’archipel. Deux études phylogénétiques récentes obtenues respectivement à partir de données moléculaires et morphologiques, ont démontré la polyphylie de Bikkia s.l., ainsi que l’existence de deux lignées évolutives : les Bikkia côtiers du Pacifi que ouest (incluant l’espèce type du genre, B. tetrandra (L.f.) A.Rich.) et les Bikkia endémiques néo-calédoniens. Il convient désormais de rétablir le nom de genre Th iollierea Montrouz., le plus ancien des noms valides et disponibles, pour les MOTS CLÉS Rubiaceae, espèces du second groupe. Des descriptions formelle et diagnostique sont ainsi Bikkia, proposées pour caractériser le genre Th iollierea et le distinguer du genre Bikkia Th iollierea, Nouvelle-Calédonie, s.s. Sept nouvelles combinaisons sont eff ectuées (T. kaalaensis (N.Hallé & Jéré- genre endémique, mie) Barrabé & Mouly, T. lenormandii (N.Hallé & Jérémie) Barrabé & Mouly, polyphylie, T. neriifolia (Brongn.) Barrabé & Mouly, T. pachyphylla (Guillaumin) Barrabé & rétablissement taxonomique, Mouly, T. parvifl ora (Schltr. & K.Krause) Barrabé & Mouly, T. retusifl ora (Brongn.) combinaisons nouvelles. Barrabé & Mouly et T. tubifl ora (Brongn.) Barrabé & Mouly). INTRODUCTION by Linnaeus fi l. for Portlandia tetrandra, making B. grandifl ora a superfl uous name (McNeill et al. Th e genus Bikkia Reinw. as currently circumscribed 2006). Five years later, Richard (1830) transferred comprises c. 20 species and occurs throughout the the Linnaean entity to Bikkia in order to distinguish Western Pacifi c, with half its members found in it from the Caribbean taxon recognized by Browne New Caledonia and the remainder shared among (Richard 1830). Some subsequent 19th century Micronesia, Melanesia, Polynesia and Indonesia. Th e authors such as Montrouzier (1860) and Brongni- type species, B. tetrandra (L.f.) A.Rich., was based art (1865) distinguished two morphological groups on a specimen from the Savage Islands (= Niue) within Bikkia. Th e fi rst of these includes coastal originally described as Portlandia tetrandra L.f. species with a broad distribution whose members (Linnaeus 1782), expanding the range of this genus, are restricted to littoral sites on limestone. Various which was initially based on material from Jamaica authors have recognized from three to ten species in (Browne 1756). Reinwardt’s (1825) protologue of this group, whose taxonomic history has been very Bikkia included a single species, B. grandifl ora Reinw. complex. Rafi nesque-Schmaltz (1820) fi rst used the ex Blume, whose type is the same specimen used name Cormigonus Raf. for the entity now known as 116 ADANSONIA, sér. 3 • 2011 • 33 (1) Reinstatement of the New Caledonian genus Th iollierea Montrouz. Bikkia, but he did not associate any species with it. sampled by Motley et al. (2005: 317, appendix) (viz. Th e later and more widely used name Bikkia was B. artensis (Montrouz.) Guillaumin, B. macrophylla recently accepted for conservation against Cormigonus (Brongn.) K.Schum., B. neriifolia (Brongn.) Schltr. (McNeill et al. 2006). Another generic name, Bik- and B. tubifl ora (Brongn.) Schltr.), along with a single kiopsis Brongn., described in 1865 for a single New sampled species of Morierina Vieill. Th e latter taxon Caledonian species (Bikkio psis pancheri Brongn., a is a genus of two species likewise endemic to New member of the coastal group), was placed in syn- Caledonia (Vieillard 1865; Brongniart & Gris 1871; onymy under Bikkia by some authors (Guillaumin Guillaumin 1948). Th e clade comprising the New 1909; Jérémie & Hallé 1976) but retained by others Caledonian endemic Bikkia and Morierina in turn (Brongniart 1865; Baumann-Bodenheim 1989). is sister to the monospecifi c genus Siemensia Urb., Th e second group of species currently included endemic to Cuba. in Bikkia comprises ten species endemic to New A second phylogenetic study based on morpho- Caledonia, all restricted to ultramafi c substrates (Jaf- logical data (Barrabé 2006), which included all fré et al. 1994) and occurring at various elevations, endemic New Caledonian members of the genus, from lowland sites to the highest mountains. Several likewise revealed two monophyletic groups that are generic names have been published to accommodate fully congruent with those identifi ed by Motley et al. members of this second group: Th iollierea Mon- (2005). Th e morphological phylogeny placed the ten trouz., initially based on a single species, T. artensis endemic New Caledonian species within a clade that Montrouz. (Montrouzier 1860), to which two more is sister to a clade comprising representatives of the species were added later (Baumann-Bodenheim previously recognized tribe Chiococceae, and these 1989); Tatea Seem. for another species (Seemann two clades are in turn sister to the genus Morierina. 1865-1873); and Grisia Brongn. for eight species Th e coastal species of Bikkia, on the other hand, (Brongniart 1865; Brongniart & Gris 1871), all form a monophyletic group, although its position of which were subsequently included in Bikkia was unresolved. Th e phylogenetic analyses of Motley by various authors (Schumann 1891; Schlechter et al. (2005) and Barrabé (2006) both suggest that 1905, 1906; Guillaumin 1909, 1930; Guillau- in order to maintain monophyly at the generic level, min & Beauvisage 1913; Jérémie & Hallé 1976). the views of earlier workers (Montrouzier 1860; A total of 17 names is validly published for the 10 Brongniart 1865; Seemann 1865-1873; Brongniart & currently recognized species of Bikkia restricted to Gris 1871) may have to be re-adopted by excluding New Caledonia (Jérémie & Hallé 1976). the New Caledonian endemics from Bikkia. As the In a molecular study of the Catesbaeeae-Chiococceae type of Bikkia (B. tetrandra) belongs to the coastal Complex (CCC) based on nuclear and chloroplast Bikkia clade, which thus comprises Bikkia s.s., it DNA data (trnL-F and ITS), Motley et al. (2005) appears necessary to place the members of the New sequenced seven of the 20 species of Bikkia recognized Caledonian endemic clade in a separate genus, for by Jérémie & Hallé (1976). While their analyses did which several names are available, viz. Grisia, Tatea, not fully resolve relationships within the complex, the and Th iollierea, the latter of which, described by results suggest that Bikkia, as currently circumscribed, Montrouzier in 1860, has priority. is not monophyletic, with members belonging to two In this paper we 1) review
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