Nomenclatural Changes in Preparation for a World Rubiaceae Checklist

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Nomenclatural changes in preparation for a World Rubiaceae Checklist

MARKUS RUHSAM1, RAFAEL GOVAERTS2 and AARON P. DAVIS FLS2* Downloaded from https://academic.oup.com/botlinnean/article/157/1/115/2418227 by guest on 23 September 2021 1University of Edinburgh, Institute of Evolutionary Biology, Ashworth Laboratories, West Main Road, Edinburgh EH9 3JT, UK 2The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, UK

Received 15 February 2006; accepted for publication 20 November 2007

During the preparation of a World Rubiaceae Checklist, numerous unplaced taxa were encountered, including illegitimate and invalid names, and species for which generic placement is uncertain. In this contribution, 35 new combinations and 20 new names are proposed, and the names of three taxa are validated. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 115–124.

ADDITIONAL KEYWORDS: Chassalia – Coptophyllum – Cyclophyllum – Diplospora – Discospermum – Global Strategy for Plant Conservation (CBD/ UNEP) – Greenea – Gynochthodes – Hoffmannia – Kohautia – Mussaenda – Mycetia – Nostolachma – Ophiorrhiza – Oxyceros – Plectronia – Pleiocarpidia – Psychotria – Psydrax – Rubiaceae – Saprosma – Tarenna – Timonius.

INTRODUCTION taxon that is invalid or illegitimate, or valid but that cannot be assigned to a known species or genus at the A widely accessible working list of known plant present time. Some of these names need to be placed species is considered to be a fundamental require- into synonymy, but others should be accepted names ment for plant conservation, and is the number one after nomenclatural attention. In this contribution, target of the Global Strategy for Plant Conservation some of the more obvious nomenclatural problems (for (Secretariat of the Convention on Biological Diversity, species that we consider should be accepted) are 2002). As part of the effort towards compiling this resolved, following the examination of type speci- working list, a complete checklist for the Rubiaceae, mens, representative herbarium material, proto- the World Rubiaceae Checklist, has been produced, logues, and other relevant literature held at the Royal including c. 615 genera and c. 13 200 accepted species Botanic Gardens, Kew (K) and the Nationaal Her- names with their synonyms: a total of c. 35 000 barium Nederland, Leiden (L). It should be empha- names. sized that no attempt has been made to sort out During the compilation, verification, and editing of problem groups and genera that are in desperate the World Rubiaceae Checklist (http://www. need of revision, or which require detailed systematic rbgkew.org.uk/wcsp/rubiaceae; Govaerts et al. in study. prep), it became evident that numerous nomencla- In this contribution, 35 new combinations and 20 tural and taxonomic problems had to be addressed in new names (nomina nova) are proposed, and the order to bring Rubiaceae nomenclature in-line with names of three taxa are validated. the current taxonomic knowledge of the family. One of the most important issues was the number of unplaced names, which, on first inspection, was MATERIAL AND METHODS around 1200. An unplaced name can be defined as any The nomenclatural types and representative herbarium material of taxa requiring new combinations *Corresponding author. E-mail: [email protected] and new names were examined using material at the

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 115–124 115 116 M. RUHSAM ET AL. herbaria K and L (herbarium abbreviations follow was asked to vote on this issue (R. K. Brummitt, Royal Holmgren, Holmgren & Barnett, 1990). In cases in Botanic Gardens, Kew, pers. comm.): P. multinervia which we were unable to examine a type or type Ridl. Journ. As. Soc. Mal. 1: 70 (1923) (distribution: specimens, new combinations and new names are Sumatera) vs. P. multinervis De Wild., Plantae proposed only for those taxa in which the protologue Bequaertianae 2: 391 (1924) (distribution: Guinea, could be used to unambiguously assign names to the Mali, Liberia), and P. brachybotrya Müll. Arg., Flora correct genera; most of these cases include those for Braziliensis 6(5): 327 (1881) (distribution: from Belize which we have provided a nomen novum for illegiti- to Brazil) vs. P. brachybotrys Ridl., Journ. Fed. Mal. mate names. Citation of the type, or types, is under- States Mus. 4: 38 (1909) (distribution: Malayan taken in the following way. If the type citation has Peninsula). The Committee, consisting of 18 members, been given in the protologue, it is given here in full; voted that the later published names are to be treated Downloaded from https://academic.oup.com/botlinnean/article/157/1/115/2418227 by guest on 23 September 2021 if we have seen the type, an exclamation mark is as homonyms. Psychotria multinervis De Wild. and given after the herbarium code (e.g. K!); it follows P. brachybotrys Ridl. are therefore illegitimate names that, if there is no exclamation mark, we have not (later homonyms). In the case of P. multinervis De seen the type. Type citations without a herbarium Wild., P. maliensis Schnell, Mém. Inst. Franç. Afrique code (after Holmgren et al., 1990) indicate that we do Noire 50: 76 (1957) becomes the accepted name, as it is not know the location of the type(s), and have thus the earliest legitimately published name currently not seen any material. considered to be a synonym of P. multinervis De Wild. (Lebrun & Stork, 1997: 204). However, a nomen novum is required for P. brachybotrys Ridl. RESULTS NOMENCLATURAL CHANGES IN PSYCHOTRIA Psychotria kerrii Govaerts, nom. nov. NOMINA NOVA AND NEW COMBINATIONS Psychotria viburnifolia Craib, Bull. Misc. Inform. Kew 1932: 483 (1932), non Psychotria viburnifolia Rusby, The genus Psychotria L. is rather easily character- Mem. New York Bot. Gard. 7: 378 (1927). ized, particularly on the basis of the inflorescence, Type: Thailand, Adang, 400 m, Kerr 14659 (K!). pyrene, and floral morphology (Petit, 1964; Robbre- cht, 1989; Davis et al., 2001; Sohmer & Davis, in prep) and sometimes on the basis of leaf characteristics. Psychotria calorhamnus (Baill.) Ruhsam, The following new combinations and new names were comb. nov. based on the examination of pyrenes and/or flowers Basionym: Uragoga calorhamnus Baill., Adansonia present on the type material, or by critical reference 12: 286 (1879). to protologues. Syntypes: New Caledonia: Poume, vi.1871, Balansa Taxonomic work on Asian Psychotria falls behind 3193 (K!); Tauté and Puebo, Deplanche 398 (P). that of the New World and Africa, and hence there are many names that require nomenclatural changes in Psychotria gaboonensis Ruhsam, nom. nov. Asia. Some taxa are still placed in the genus Uragoga Cephaelis latifolia Hiern in D.Oliver, Fl. Trop. Afr. 3: Baill., but this is a synonym of Psychotria (Petit, 226 (1877), non Psychotria latifolia Humb. & Bonpl. 1964). The genus Cephaelis has been shown to be an ex Schult. in J.J. Roemer & J.A. Schultes, Syst. Veg. artificial concept, consisting of several genera (e.g. 5: 189 (1819). Andersson, 2002: 194), with most species occurring in Syntypes: West Africa, Gaboon River [Equatorial Asia now placed in synonymy with either Psychotria Guinea/Gabon], Mann 931 (K!); loc. cit., Mann 939 or Chassalia Comm. ex Poir. (see ‘New combinations (K!). and nomina nova in Chassalia’, below). Cephaelis is not recognized as a separate genus, and is a generic Psychotria kumbangii Ruhsam, nom. nov. synonym of Psychotria. Psychotria patula Ridl., J. Fed. Malay States Mus. 8(4): 51 (1917), non Psychotria patula Humb. & Psychotria tahanensis Ruhsam, nom. nov. Bonpl. ex Schult. in J.J.Roemer & J.A.Schultes, Syst. Psychotria brachybotrys Ridl., Journ. Fed. Mal. States Veg. 5: 189 (1819). Mus. 4: 38 (1909), non Psychotria brachybotrya Müll. Type: Sumatra, Sungei Kumbang, 4000–5000 ft Arg., Flora Braziliensis 6(5): 327 (1881). [1120–1400 m], Robinson & Kloss s.n. (K!). Type: No type cited in the protologue, only the locality ‘Gunong Tahan’. Psychotria matogrossensis Govaerts, nom. nov. Notes: To establish whether the following names in Cephaelis krauseana Standl., Publ. Field Mus. Nat. Psychotria are to be considered homonyms, the Hist., Bot. Ser. 8: 181 (1930), non Psychotria krau- Committee for the Nomenclature of Vascular Plants seana Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 8:

212 (1930); Psychotria krauseana (Standl.) Zappi, Psychotria ligustrina (Baill.) Guillaumin, Arch. Bot., Prodr. Fl. Matogross., Ser. B, 3: 258 (1998), nom. illeg. Caen, 3. Mem. No. 5: 31 (1930), nom. illeg. Type: Brazil, Matto Grosso, Rio Arnos, xii.1914, Kuhl- Type: New Caledonia, Tchiaor and Puebo, Balansa mann 1410. 3194 (P). Notes: Psychotria krauseana (Standl.) Zappi was Notes: Psychotria ligustrina (Baill.) Guillaumin was made as a new combination for Cephaelis krauseana made as a new combination for Uragoga ligustrina Standl., but is a later homonym of Psychotria krau- Baill., but is a later homonym of Psychotria ligustrina seana Standl. A new name is therefore required. Willd. ex Schult. A new name is therefore required.

Psychotria melintangensis Govaerts, nom. nov. Psychotria srilankensis Ruhsam, nom. nov. Downloaded from https://academic.oup.com/botlinnean/article/157/1/115/2418227 by guest on 23 September 2021 Psychotria coriacea Korth., Ned. Kruidk. Arch. 2: 242 Grumilea stenophylla Thwaites, Enum. Pl. Zeyl.: 147 (1851) non Psychotria coriacea Poir. ex Spreng., Syst. (1859), non Psychotria stenophylla Spreng., Syst. Veg. 1: 742 (1824). Veg. 1: 744 (1824); Psychotria stenophylla (Thwaites) Type: No type cited in the protologue, only two locali- Hook.f., Fl. Brit. Ind. 3: 161 (1880), nom. illeg. ties: Melintang (Sumatera) and Balaron (Borneo). Type: Sri Lanka, 1853, Thwaites C. P. 2609 (K!). Notes: Psychotria stenophylla (Thwaites) Hook.f. was Psychotria mons-mi Ruhsam, nom. nov. made as a new combination for Grumilea stenophylla Uragoga balansae Baill., Adansonia, 12: 223 (1879) Thwaites, but is a later homonym of Psychotria steno- non Psychotria balansae Pitard in Lecomte, Fl. Indo- phylla Spreng. A new name is therefore required. Chine 3: 353 (1924); Psychotria balansae (Baill.) Guil- laumin, Arch. Bot., Caen, 3. Mem. No. 5: 27 (1930), nom. illeg. Psychotria thailandensis Ruhsam, nom. nov. Syntypes: New Caldeonia: Mt. Mi, 1000 m, Balansa Psychotria polita Craib, Bull. Misc. Inform. Kew 1120 (P); Mt. Arago, 1200 m, Balansa 3424 (P). 1932: 482 (1932), non Psychotria polita Valeton, Bot. Notes: Psychotria balansae (Baill.) Guillaumin was Jahrb. Syst. 61: 95 (1927). made as a new combination for Uragoga balansae Type: Thailand, Ranawng, La-un, 20 m, Kerr 16510 Baill., but is a later homonym of Psychotria balansae (K!). Pitard. A new name is therefore required. Psychotria vietnamensis Ruhsam, nom. nov. Psychotria namwingensis Govaerts, nom. nov. Chassalia ovoidea Pierre ex Pit. in H.Lecomte, Fl. Psychotria alata Craib, Bull. Misc. Inform. Kew 1932: Indo-Chine 3: 366 (1924), non Psychotria ovoidea 476 (1932) non Psychotria alata Valeton, Bot. Jahrb. Wall. ex Hook.f., Fl. Brit. India 3: 166 (1880); Syst. 61: 80 (1927). Psychotria ovoidea (Pierre ex Pit.) P.H. Ho, Cayco Type: Thailand, Surat, Klawng Nam Wing, 100 m, Vietnam 3(1): 246 (1993), nom. illeg. Kerr 12246. Syntypes: Cochinchine, Thorel s.n. [79S?] (K!); loc. cit. Harmand s.n., loc. cit. Talmy s.n., loc. cit. Pierre s.n., Psychotria oblanceolata (R.D. Good) Ruhsam, loc. cit. Godefrey s.n., loc. cit. Poilane s.n. comb. nov. Notes: The pyrene morphology of the specimen Thorel Basionym: Uragoga oblanceolata R.D. Good, J. Bot. s.n. clearly shows that Chassalia ovoidea Pierre ex 64 (Suppl. 2): 36 (1926). Pit. should be transferred to Psychotria. Psychotria Type: Angola, in the M’Bulu hills, R. N’Zaza, R. Lufo, ovoidea (Pierre ex Pit.) P.H. Ho was made as a new Gossweiler 7760 (K!). combination for Chassalia ovoidea Pierre ex Pit., but this is a later homonym of Psychotria ovoidea Wall. ex Psychotria poissoniana (Baill.) A.P. Davis, Hook.f. A new name is therefore required. comb. nov. Basionym: Uragoga poissoniana Baill., Adansonia Psychotria gjellerupii A.P. Davis, nom. nov. 12: 230 (1879). Cephaelis papuana Valeton, Nova Guinea 8: 767 Syntypes: New Caledonia: above Conception, 700 m, (1912), non Psychotria papuana (Wernham) H.St. 9.xi.1868, Balansa 340 (K!); loc. cit., Balansa 1119 (P); John, Occas. Pap. Bernice Pauahi Bishop Mus. 12(7): loc. cit., Balansa 2028 (P); loc. cit., Balansa 2028a (P). 3 (1936). Type: New Guinea, Begowri [Begoure] River, Psychotria pueboensis Ruhsam, nom. nov. 24.vi.1910, Gjellerup 205 [(‘483’)] (L!). Uragoga ligustrina Baill., Adansonia 12: 253 (1879), Notes: Valeton (1912) no doubt placed this taxon in non Psychotria ligustrina Willd. ex Schult. in J.J. Cephaelis Sw. on account of its condensed head-like Roemer & J.A. Schultes, Syst. Veg. 5: 188 (1819); inﬂorescences. The specimen Gjellerup 205 shows the

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 115–124 118 M. RUHSAM ET AL. very typical Psychotria features of leaves drying Chassalia griffithii (Hook.f.) A.P. Davis, pinkish-brown, and short, straight corolla tubes. comb. nov. Basionym: Cephaelis griffithii Hook.f., Fl. Brit. India Psychotria warburgiana A.P. Davis, nom. nov. 3: 178 (1880). Psychotria apiculata Warb., Bot. Jahrb. Syst. 13: 439 Synonym: Psychotria malaccensis I.M.Turner, Asian (1891), non Psychotria apiculata Müll.Arg., Flora 59: J. Trop. Biol., 1(2): 27 (1995). 552 (1876). Syntypes: Malay Peninsula, Malacca, Griffith s.n. Type: New Guinea, Sattelburg, iii.–iv.1869, Warburg (Kew Distr. 3032) (K!); loc. cit. Maingay 928. 21452 [lectotype BM (designated by Sohmer, 1988: 29)]. Downloaded from https://academic.oup.com/botlinnean/article/157/1/115/2418227 by guest on 23 September 2021 Notes: We have not examined any original material of Chassalia ridleyi (King) A.P. Davis, comb. nov. P. apiculata, although there is a photograph of the Basionym: Cephaelis ridleyi King, J. Asiat. Soc. lectotype in Sohmer (1988: 20, fig. 3). Bengal, Pt. 2, Nat. Hist. 73: 106 (1904). Synonym: Psychotria naviculiformis I.M.Turner, Asian J. Trop. Biol., 1(2): 27 (1995). NEW COMBINATIONS AND NOMINA NOVA Type: Malay Peninsula, Selangor, Bukit Kutu, 3000 ft IN CHASSALIA [914 m], v.1896, Ridley 7417 (K!). In a similar way to Psychotria, the genus Chassalia is rather easily characterized on the basis of salient Chassalia melanocarpa (Ridl.) A.P. Davis, morphological features (for example, Piesschaert, Rob- comb. nov. brecht & Smets, 2001: 121). The stipules and stipule Basionym: Cephaelis melanocarpa Ridl., J. Straits remains (basal parts) of Chassalia are nearly always Branch Roy. Asiat. Soc. 86: 298 (1922). characteristically ‘chaffy’ (membranaceous, whitish, Synonym: Psychotria albopedunculata I.M.Turner, and rather corky) when dry; the inflorescences are Asian J. Trop. Biol., 1(2): 27 (1995). always terminal, with often rather fleshy or succulent, Type: Malay Peninsula, Negri Sembilan, Bukit and sometimes brightly coloured (e.g. red, purple, white), inflorescence parts (for example, branches, Tangga, 2500 ft [762 m], xii.1920, Ridley s.n. (K!). pedicels, bracts); the corolla tubes are nearly always long and slightly to markedly curved; the fruits are Chassalia angustifolia (Ridl.) A.P. Davis, more or less smooth when dry, although some species comb. nov. have pyrenes with a dorsal ridge which shows as a Basionym: Cephaelis angustifolia Ridl., J. Straits ridge or line on the dry fruit; the pyrenes mostly have Branch Roy. Asiat. Soc. 79: 88 (1918). a distinct ventral excavation, usually a short pre- Synonym: Psychotria ophirensis I.M.Turner, Asian J. formed germination slit (PGS) on the dorsal surface (if Trop. Biol., 1(2): 26 (1995). a dorsal ridge is present, this running along the ridge), Syntypes: Malay Peninsula: Johor, Gunong Pantai, and usually two short lateral PGSs (Piesschaert et al., 7.xii.1892, Ridley s.n. (K!); Malacca, Mt. Ophir, 1999, 2001; Piesschaert, 2001). In addition, Chassalia Maingay s.n.; Malacca, Mt. Ophir, Griffith s.n.; Negri shoot bark is normally dark brown to black, the leaves Sembilan, Gunong Tampin [no collector cited]. of most species are membranaceous, the corolla buds are often slightly winged, and the pollen grains are large, colpate or porate, with a reticulate exine (Piess- Chassalia tricepa (Ridl.) A.P. Davis, comb. nov. chaert et al., 2001). The following new combinations Basionym: Cephaelis tricepa Ridl., J. Straits Branch and new names were based on the examination of Roy. Asiat. Soc. 79: 89 (1918). vegetative features (i.e. stipules, shoot bark, leaves), Synonym: Psychotria tricepa (Ridl.) I.M.Turner, pyrenes, and/or flowers, as found on type material and Asian J. Trop. Biol., 1(2): 27 (1995). representative specimens. Type: Malay Peninsula, Selangor, Sempang mines, In this section, we place a number of species origi- Semangkok Pass, 1911, Ridley 15664 (K!). nally placed in Cephaelis into Chassalia. As stated above, Cephaelis is a synonym of Psychotria, with most species from Asia now being placed in either Chassalia Chassalia porcata (I.M.Turner) A.P. Davis, or Psychotria. Turner (1995) transferred several Ceph- comb. nov. aelis species to Psychotria, but, on revisiting the type Basionym: Psychotria porcata I.M.Turner, Asian J. specimens, we have found that they have the salient Trop. Biol., 1(2): 27 (1995). characteristics of Chassalia (see above) and not Psy- Synonym: Cephaelis elongata Ridl., J. Straits Branch chotria. The necessary nomenclatural alterations, Roy. Asiat. Soc. 79: 90 (1918), non Chassalia elongata amongst our other changes, are provided below. Hutchinson & Dalziel, Fl. W. Trop. Afr. 2: 127 (1931).

Type: Malay Peninsula, Selangor, Semangkok Pass, Type: Java, Pasir Kempuk, 1100 m, 1913, Backer 1911, Ridley s.n. (K!). 11049 (BO, L!).

Chassalia subcoriacea (Ridl.) A.P. Davis, NEW COMBINATIONS IN VANGUERIEAE comb. nov. There are three genera of Vanguerieae Dumort. in Basionym: Cephaelis subcoriacea Ridl., J. Fed. Malay south and south-east Asia: Canthium Lam., Psydrax States Mus. 5: 36 (1914). Gaertn., and Cyclophyllum Hook.f. The informal Synonym: Psychotria subcoriacea (Ridl.) I.M.Turner, Pyrostria ‘group B’ was recognized by Bridson Asian J. Trop. Biol., 1(2): 27 (1995). (1987), and represents a group of taxa that may Type: Malay Peninsula, Selangor, Gunong Downloaded from https://academic.oup.com/botlinnean/article/157/1/115/2418227 by guest on 23 September 2021 either constitute a new genus or perhaps a disjunct Mengkuang Lebah, 16.i.1913, Robinson s.n. (K!). part of Pyrostria (Bridson, 1987). It is likely, then, that there are four genera of Vanguerieae in south Chassalia tahanica (I.M.Turner) A.P. Davis, and south-east Asia. Canthium, Psydrax, Cyclophyl- comb. nov. lum, and Pyrostria ‘group B’ are easily distinguished Basionym: Psychotria tahanica I.M.Turner, Asian J. on morphological grounds (for example, Bridson, Trop. Biol., 1(2): 26 (1995). 1987; Reynolds & Henderson, 1999: 354); molecular Synonym: Cephaelis albiflora Ridl., J. Fed. Malay data (Lantz & Bremer, 2004) support the mainte- States Mus. 6: 155 (1915), non Chassalia albiflora nance of Cyclophyllum, Psydrax, and Pyrostria sensu Krause, Bot. Jahrb. Syst. 39: 566 (1907). stricto. Lantz & Bremer (2004: 279) restrict Type: Malay Peninsula, Pahang, Gunong Tahan, on Canthium to a group of mostly spiny stemmed the Padang, 5000 ft [1524 m], vii.1911, Ridley 16075 plants, which we interpret as having a distribution (K!). from Africa to southern Asia (including India and Sri Lanka), but excluding Madagascar and south- east Asia. Lantz & Bremer (2004) clearly show that Chassalia singapurensis (Ridl.) A.P. Davis, Canthium is polyphyletic as currently circumscribed. comb. nov. Similarly, Plectronia L. was formerly used in Rubi- Basionym: Cephaelis singapurensis Ridl. J. Straits aceae as the generic name for a large number of Branch Roy. Asiat. Soc. 79: 89 (1918); Psychotria unrelated species, mostly belonging to the tribe singapurensis (Ridl.) I.M.Turner, Asian J. Trop. Biol., Vanguerieae. However, Plectronia must be restricted 1(2): 27 (1995). to the family Oliniaceae, as Plectronia is a generic Type: Singapore, Bajau, 1892, Ridley 4966 (K!). synonym of Olinia Thunb. (Ross, 1975: 491; Verdcourt, 1987: 127). Chassalia elliptica (Ridl.) A.P. Davis, comb. nov. Cyclophyllum is restricted to south-east Asia, Aus- Basionym: Cephaelis elliptica Ridl., J. Straits Branch tralasia, and the Pacific (Bridson, 1987: 614, map 1). Roy. Asiat. Soc. 79: 89 (1918). Psydrax is found throughout the Old World, from Type: Malay Peninsula, Pahang, Telom, vi.1908, West Africa to the Pacific. Canthium is probably Ridley 13636 (K!). restricted to Africa and southern Asia (see above), and Pyrostria ‘group B’ occurs from mainland south-east Asia to the Kepala Burung (Vogelkop Peninsula) in Chassalia blumeana Govaerts, nom. nov. the extreme north-west part of New Guinea (see Cephaelis stipulacea Blume, Bijdr: 1005 (1826), non Bridson, 1987: 614, map 1). Chassalia stipulacea DC., Prodr. 4: 532 (1830); Chas- It is proposed that four Rubiaceae species currently salia stipulacea (Blume) Piessch., Flora (Germany) placed in Plectronia should be transferred to Cyclo- 196: 128 (2001), nom. illeg. phyllum. The type specimens of Plectronia marque- Type: No type cited, just two locations: Seribu, Salak. sensis F.Br., Pl. sessilifolia Valeton, Pl. subsessilis Notes: Chassalia stipulacea (Blume) Piessch. was Valeton, and Pl. urophylla Valeton show the salient made as a new combination for Cephaelis stipulacea characteristics of Cyclophyllum after Bridson (1987: Blume, but is a later homonym of Chassalia stipula- 616) (see also Davis & Ruhsam, 2005). In addition, it cea DC. [Prodr. 4: 532 (1830)]. A new name is there- is considered that the following three species cur- fore required. rently placed in Canthium, C. calcicola Craib, C. fasciculatum Blume, and C. longistylum Merr., and one Chassalia nannochlamys (Bakh.f) A.P. Davis, in Plectronia, P. amplifolia Elmer, belong in the genus comb. nov. Psydrax. The type specimens and representative Basionym: Cephaelis nannochlamys Bakh.f., Blumea material of these four species possess the critical 12: 64 (1963). diagnostic features of Psydrax (e.g. Bridson, 1998:

345). The following new combinations and new names Type: New Guinea, Sepik, Lordberg, 1000 m, are provided. 4.xii.1912, Ledermann 10045 [a] (L!).

Cyclophyllum subsessile (Valeton) A.P. Davis, NEW COMBINATIONS IN PSYDRAX comb. nov. Psydrax calcicola (Craib) A.P. Davis, comb. nov. Basionym: Plectronia subsessilis Valeton, Bot. Jahrb. Basionym: Canthium calcicola Craib, Fl. Siam. 2: Syst. 61: 58 (1927). 136 (1932). Type: New-Guinea, Sepik, Etappenberg, 850 m, Type: Thailand, Nakawn Sritamarat, Patalung, Kao 17.x.1912, Ledermann 9347 (L!). Hua Trek, 50 m, 2.v.1930, Kerr 19294 (K!). Downloaded from https://academic.oup.com/botlinnean/article/157/1/115/2418227 by guest on 23 September 2021 Cyclophyllum urophyllum (Valeton) A.P. Davis, comb. nov. Psydrax fasciculata (Blume) A.P. Davis, Basionym: Plectronia urophylla Valeton, Bot. Jahrb. comb. nov. Syst. 61: 58 (1927). Basionym: Canthium fasciculatum Blume, Bijdr.: Syntypes: New Guinea: Jaduna, 300 m, 19.iv.1909, 967 (1826). Schlechter 19277 (L!); Somadjidji, 450 m, 5.v.1909, Type: Java, Blume s.n. (K!). Schlechter 19353 (L!).

Psydrax longistyla (Merr.) A.P. Davis, comb. nov. MISCELLANEOUS NOMENCLATURAL CHANGES Basionym: Canthium longistylum Merr., Pap. Michi- IN RUBIACEAE gan Acad. Sci. 23: 195 (1937 publ. 1938). NEW NAMES FOR OPHIORRHIZA AND MUSSAENDA Type: Sumatra, Laboehan Batoe, Kota Pinang, Langga Pajoeng (on the Soengei Kanan), The following two names are illegitimate and require 7–30.iii.1933, Rahmat 3601 (K!). new combinations.

Ophiorrhiza pinatuboensis Ruhsam, nom. nov. Psydrax amplifolia (Merr.) A.P. Davis, comb. nov. Ophiorrhiza bicolor Elmer, Leafl. Philipp. Bot. 9: 3211 Basionym: Plectronia amplifolia Elmer, Leafl. (1934), non Ophiorrhiza bicolor Craib, Fl. Siam. 2: 62 Philipp. Bot. 9: 3261 (1934). (1932). Type: Philippines, Island of Luzon, Province of Sor- Type: Philippines, Luzon, Pampanga Province, Mt. sogon, Irosin (Mt. Bulusan), vii.1916, Elmer 16739 Pinatubo, v.1927, Elmer 21958 (K!). (L!). Mussaenda sandakana Govaerts, nom. nov. Psydrax sepikensis A.P. Davis, nom. nov. Mussaenda laxiflora Merr., Univ. Calif. Publ. Bot. 15: Plectronia grandifolia Valeton, Bot. Jahrb. Syst. 61: 280 (1929), non Mussaenda laxiflora Hutch. in C.S. 55 (1927), non Psydrax grandifolia (Thwaites) Rids- Sargent, Pl. Wilson. 3: 399 (1916). dale, Blumea, 41: 462 (1996). Type: British North Borneo [Sabah], Myburgh Prov- Syntypes: New-Guinea, Sepik, 20–50 m: Ledermann ince, Sandakan, x.–xii.1921, Elmer 20282 (K!). 6998, 12.iv.1912 (L!); loc. cit. Ledermann 10508, 9.i.1913 (L!). NEW COMBINATIONS IN TIMONIUS, HOFFMANNIA, MYCETIA, GREENEA, PLEIOCARPIDIA, GYNOCHTHODES, SAPROSMA, DIPLOSPORA, NEW COMBINATIONS IN CYCLOPHYLLUM DISCOSPERMUM, COPTOPHYLLUM, AND TARENNA Cyclophyllum marquesense (F.Br.) Govaerts, Timonius uniflorus (C.F. Gaertn.) Govaerts, comb. nov. comb. nov. Basionym: Plectronia marquesensis F.Br., Bull. Basionym: Erithalis uniflora C.F. Gaertn., Suppl. Bishop Mus., Honolulu, 130: 293 (1935). Carp. 92 (tab. 196) (1806). Type: Marquesas, Hivaoa, Atuona, 8.xii.1921, Brown Type: No type cited. 826 (K!). Synonyms: Erithalis polygama G.Forst., Fl. Ins. Austr: 17 (1786). Erithalis cymosa G.Forst. ex Spreng. Cyclophyllum lordbergense A.P. Davis, nom. nov. Pl. Min. Cogn. Pug. 1: 17 (1813). Burneya forsteri Plectronia sessilifolia Valeton, Bot. Jahrb. Syst. 61: Cham. & Schltdl., Linnaea 4: 189 (1829), nom. 57 (1927), non Cyclophyllum sessilifolium (A.Gray) superfl. Timonius forsteri DC., Prodr. 4: 461 (1830), A.C.Sm. & S.P.Darwin, Fl. Vitiensis Nova, 4: 236 nom. superfl. Polyphragmon minus A.Rich. ex DC., (1988). Prodr. 4: 445 (1830), nom. superfl. Erithalis cymosa

G.Forst. ex DC., Prodr. 4: 465 (1830), pro syn. Bobea Notes: The genus Rhombospora Korth. is a synonym gaertneri Korth., Ned. Kruidk. Arch. 2(2): 211 (1851), of Greenea Wight & Arn. (Brummitt, 1992). Type nom. superfl. Bobea forsteri (DC.) Korth., Ned. material of Rhombospora commersonii Korth., held at Kruidk. Arch. 2(2): 211 (1851), nom. superfl. Timonius Kew and annotated by C. Tange, confirms that a new polygamus (G.Forst.) Rob., Proc. Amer. Acad. Arts 45: combination is needed. 408 (1910), later homonym. Notes: Although Timonius polygamus (G.Forst.) Rob. Pleiocarpidia maingayi (King & Gamble) is an accepted name in Flora Vitensis Nova (Smith, Ruhsam, comb. nov. 1988: 158), it is actually a later homonym of T. po- Basionym: Aulacodiscus maingayi King & Gamble, lygamus Oken [Allg. Naturgesch. 3(2): 1039 (1841)] J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 72: 189 (1903). and therefore illegitimate. The basionym of Timonius Syntypes: Malay Peninsula: Malacca, Maingay 938 Downloaded from https://academic.oup.com/botlinnean/article/157/1/115/2418227 by guest on 23 September 2021 polygamus (G.Forst.) Rob. is Erithalis polygama (K!); Selangor, Curtis 2343; Perak, Scoretechini 2004; G.Forst., which has a host of illegitimate nomina loc. cit., King’s Collector 1086. nova (R. Govaerts, unpubl. data). The earliest avail- Notes: The genus Aulacodiscus Hook.f. is a synonym able legitimately published synonym for replacing of Pleiocarpidia K. Schum. (Bremekamp, 1940; Brum- T. polygamus (G.Forst.) Rob. is Erithalis uniflora C.F. mitt, 1992) with one unplaced species. A new combi- Gaertn. A new combination is therefore necessary. nation is required.

Gynochthodes motleyi (Hook.f.) Ruhsam, Hoffmannia lindenii (Kuntze) A.P. Davis, comb. nov. comb. nov. Basionym: Tetralopha motleyi Hook.f., Hooker’s Icon. Basionym: Xerococcus lindenii Kuntze, Revis. Gen. Pl. 11: tab. 1072 (1870). Pl. 1: 285 (1891). Type: Borneo, Motley s.n. (K!). Type: Mexico, Chiapas, Linden 1638 (K!). Notes: See notes below. Notes: Xerococcus Oerst. is a synonym of Hoffmannia Sw. (Brummitt, 1992). All species of the former genus Gynochthodes polillensis (Hook.f.) Ruhsam, (apart from X. lindenii) have been transferred to the comb. nov. latter. Basionym: Tetralopha polillensis C.B. Robinson, Philipp. Journ. Sc., Bot. 6: 226 (1911). Type: Philippines, Polillo, about 1.5 km south of the Mycetia apoensis (Elmer) Govaerts, comb. nov. town, Robinson BS 9064. Basionym: Adenosacme apoensis Elmer, Leafl. Notes: The genus Tetralopha Hook.f. is a synonym of Philipp. Bot. 3: 1001 (1911). Gynochthodes Blume (Brummitt, 1992), with cur- Type: Philippines, Mindanao, District of Davao, rently two unplaced taxa: Tetralopha motleyi Hook.f. Todaya (Mt. Apo), v.1909, Elmer 10504 (K!). and Tetralopha polillensis C.B. Robinson. In the Notes: The genus Adenosacme Wall. is a synonym of protologue of T. polillensis, Robinson (1911: 226) is Mycetia Reinw. (for example, Deb, 1988; Gideon, uncertain about the generic placement of this species, Tjitrosoedirdjo & Veldkamp, 1988; Brummitt, 1992), and mentions that ‘the species might with almost with A. apoensis Elmer and A. mindanaensis Elmer equal propriety be placed in Gynochthodes’asit the only unplaced taxa remaining in Adenosacme. shows the salient characters of that genus. The type Two new combinations are therefore needed. specimen of T. motleyi Hook.f. also exhibits the typical characters of Gynochthodes, including unisexual Mycetia mindanaensis (Elmer) Govaerts, flowers in more or less congested cymes, short comb. nov. pedicels, calyx reduced to a ring-like rim, and calyx Basionym: Adenosacme mindanaensis Elmer, Leafl. tube pilose inside. We therefore provide two new Philipp. Bot. 3: 1002 (1911). combinations. Type: Philippines, Mindanao, District of Davao, Todaya (Mt. Apo), vi.1909, Elmer 10831 (K!). Saprosma elegans (Korth) A.P. Davis, comb. nov. Notes: See above. Basionym: Cleisocratera elegans Korth., Verh. Nat. Gesch. Ned. Bezitt., Bot.: 257, tab. 62 (1844). Type: Borneo, Korthals s.n. (L!). Greenea commersonii (Korth) Tange ex Ruhsam, Notes: The study of herbarium specimens held at L comb. nov. confirms that the genus Cleisocratera Korth. is a Basionym: Rhombospora commersonii Korth., Ned. synonym of Saprosma Blume (Brummitt, 1992). The Kruidk. Arch. 2: 114 (1850). woody fimbriate stipules, which, when they fall, leave Type: Sumatera, Korthals s.n. (K!). many fibrous/woody remains at the stipule line, is a

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 115–124 122 M. RUHSAM ET AL. good diagnostic character for Saprosma and is clearly into Pomazota. Subsequently, Pomazota species have evident in C. elegans Korth. The terminal inflores- been placed into Coptophyllum, although some cences and small four-merous, delicate, white, and species remain in the former genus. We have rather fleshy flowers are also characteristic of seen the types of two species of Pomazota, which Saprosma. We have seen numerous flowering speci- clearly have the salient characters of Coptophyllum mens of C. elegans in L, but no material bearing fruit, (Bremekamp, 1947), and therefore propose two and we have not been able to find a copy of the further combinations. original protologue (Korthals, 1844; missing herb. K). However, the description of flowers and fruit by Coptophyllum vanleeuwenii (Bremek.) A.P. Davis, Bentham & Hooker (1873: 123), who would have had comb. nov. Downloaded from https://academic.oup.com/botlinnean/article/157/1/115/2418227 by guest on 23 September 2021 access to the protologue and the figure of C. elegans Basionym: Pomazota vanleeuwenii Bremek., J. (Korthals, 1844), unambiguously matches the diag- Arnold Arbor. 28: 197 (1947). nostic features of Saprosma. Type: Sumatra, Haboko, 21.ii.1919, Doctors van Leeuwen 3276 (L!). Diplospora sorsogonensis (Elmer) A.P.Davis, comb. nov. Tarenna celebica (Miq.) Ruhsam, comb. nov. Basionym: Tricalysia sorsogonensis Elmer, Leafl. Basionym: Stylocoryna celebica Miq., Ann. Mus. Bot. Philipp. Bot. 9: 3269 (1934). Lugduno-Batavi 4: 190 (1869). Type: Philippines, Province of Sorsogon, Island of Type: Celebes, Forsten s.n. (L!). Luzon, Irosin (Mt. Bulusan), ix.1916, Elmer 17261. Notes: Stylocoryna Cav. is a synonym of Aidia Lour. Notes: Species of ‘Asian Tricalysia’ are mostly refer- (Brummitt, 1992), although most species formerly able to the genera Diplospora DC. and Discospermum described in this genus have been placed in other (Ali & Robbrecht, 1991). The smallish fruit (less than genera, most notably Tarenna Gaertn. The type speci- 1 cm wide) and rather few seeds per locule (four or mens of S. celebica and S. forsteniana represent flow- five), coupled with the Philippine location, place ering material and possess the characteristic features T. sorsogonensis most satisfactorily in Diplospora (Ali of Tarenna (for example, Bridson & Verdcourt, 1988). & Robbrecht, 1991). We have not seen the type speci- The flowers of both species are four-merous, which, men of T. sorsogonensis, but have studied another although uncommon in Tarenna, is within the circum- collection from the same locality [loc. cit., ix.1916, scription of the genus. Pavetta is the only genus likely Elmer 17164 (L)]. to be confused with Tarenna in Asia, and, indeed, the former usually has four-merous flowers. However, Discospermum polyspermum (Valeton) Ruhsam, Pavetta usually has chartaceous leaves with bacterial comb. nov. nodules, a distinctive type of branching (‘Pavetta Basionym: Diplospora polysperma Valeton, Icon. branching’), and usually two (rarely one by abortion) Bogor.: tab. 142 (1904). seeds per fruit. Stylocoryna celebica and S. forste- Type: Cult. in Hort. Bogor, IV. E. 130 (L!). niana have numerous ovules per locule, subcoriaceous Notes: The large, thick-walled fruit and numerous leaves lacking bacterial nodules, and stipules drying seeds per locule place this species into Discosper- blackish. We propose two new combinations in mum rather than Diplospora (Ali & Robbrecht, Tarenna. It seems possible that S. celebica and 1991). Ali & Robbrecht (1991: 299) did not make a S. forsteniana are conspecfic; further critical study combination for this species in Discospermum, is required. because they argued that it could have received another name in its country of origin, possibly Tarenna forsteniana (Miq.) Ruhsam, comb. nov. under a different generic name (for example, the Basionym: Stylocoryna forsteniana Miq., Ann. Mus. artificial concept for Randia). Bot. Lugduno-Batavi 4: 191 (1869). Type: Celebes, in sylvis Tondano, Forsten s.n. (L!). Coptophyllum simalurense (Bremek.) A.P. Davis, comb. nov. Basionym: Pomazota simalurensis Bremek., J. MISCELLANEOUS INVALID NAMES Arnold Arbor. 28: 197 (1947). Kohautia caespitosa ssp. amaniensis (K.Krause) Type: Sumatra, Is. Simaluoen [?], 4.vii.1918, Achmad Govaerts, comb. et stat. nov. 516 (L!). Basionym: Oldenlandia amaniensis K.Krause, Bot. Notes: Bremekamp (1947) came to the conclusion Jahrb. Syst. 43: 127 (1909). that Pomazota Ridl. and Coptophyllum Korth. are Notes: Kohautia caespitosa ssp. amaniensis congeneric, but incorrectly rejected the earlier pub- (K.Krause) Mantell ex Puff, nom. inval. [Rubiac. Fl. lished Coptophyllum, and placed his accepted names Ethiopia: 31 (1989)], was published without either a

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 115–124 NOMENCLATURAL CHANGES FOR A WORLD RUBIACEAE CHECKLIST 123 basionym reference or full and direct reference to its ACKNOWLEDGEMENTS author and place of valid publication, and is thus an We gratefully acknowledge the Global Biodiversity invalid name according to the International Code of Information Facility (GBIF), within the Electronic Botanic Nomenclature (Greuter et al., 2000: Art. 33.3). Catalogue of Names of Known Organisms (ECAT) The full basionym reference is provided here, thus programme, who supported most of the work under- validating the name Kohautia caespitosa ssp. taken for this contribution. We would also like to amaniensis. thank the Director and Staff at the Nationaal Her- barium Nederland, Leiden University branch, and R.K Brummitt (Royal Botanic Gardens, Kew) for their Oxyceros hoaensis (Pierre ex Pit.) Tirveng. ex assistance. We are also grateful to Elmar Robbrecht Downloaded from https://academic.oup.com/botlinnean/article/157/1/115/2418227 by guest on 23 September 2021 Ruhsam, comb. nov. (National Botanic Garden of Belgium, Meise) for Basionym: Randia hoaensis Pierre ex Pit. in H.Le- reviewing an earlier version of this contribution. comte, Fl. Indo-Chine 3: 235 (1923). Notes: The combination Oxyceros hoaensis (Pierre ex REFERENCES Pit.) Tirveng., nom. inval. [Nordic J. Bot. 3: 466 (1983), was published without basionym date and is Ali SJ, Robbrecht E. 1991. Remarks on the tropical Asian therefore invalid (Greuter et al., 2000: Art. 33.3). and Australian taxa included in Diplospora or Tricalysia (Rubiaceae-Ixoroideae-Gardenieae). Blumea 35: 279–305. Andersson L. 2002. Relationships and generic circumscrip- Nostolachma viridiflora (Ridl.) J.-F.Leroy ex A.P. tion in the Psychotria complex (Rubiaceae, Psychotrieae). Davis, comb. nov. Systematics and Geography of Plants 72: 167–202. Basionym: Coffea viridiflora Ridl., J. Straits Branch Bentham G, Hooker JD. 1873. Genera plantarum. Caprifo- liaceas to Compositas, Vol. 2, Part 1. London: Lovell Reeve. Roy. Asiat. Soc. 79: 82 (1918). Bremekamp CEB. 1940. A monograph of the genus Pleio- Synonym: Lachnastoma viridiflorum (Ridl.) A.Chev., carpidia K. Sch. Recueil des Travaux Botaniques Néerlan- Encycl. Biol. 22(2): 35 (1942). dais 37: 198–236. Syntypes: Malaysian Peninsula: Selangor, Batu Bremekamp CEB. 1947. A monograph of the genus Poma- Caves, 4.iii.1915, Ridley s.n. (K!); Perak, Waterloo, zota Ridley. Journal of the Arnold Arboretum 28: 186–203. xii.1887, Curtis 1304 (K!). Bridson DM. 1987. Studies in African Rubiaceae – Notes: Coffea viridiflora Ridl. was transferred to Vanguerieae: a new circumscription of Pyrostria and a new Lachnastoma Korth. by Chevalier (1942), and then to subgenus, Canthium subgen. Bullockia. Kew Bulletin 42: Nostalachma T.Durand by Leroy (Lobreau-Callen 611–639. & Leroy, 1980; see above). Nostalachma replaces Bridson DM. 1998. Rubiaceae. In: Pope GV, ed. Flora Zamb- Lachnastoma because of the earlier use of this genus esiaca, Vol. 5, Part 2. Surrey: Royal Botanic Gardens, Kew, name in the Asclepiadaceae (Lachnostoma Kunth.). 211–377. The combination Nostolachma viridiflora (Ridl.) Bridson DM, Verdcourt B. 1988. Rubiaceae. In: Polhill RM, J.-F.Leroy [Ass. Sci. Internat. Café, 9e Colloque: 481 ed. Flora of tropical East Africa, Rubiaceae, Part 2. Rotter- (1980)] was published without citation of the dam: Balkema, 415–747. basionym volume and the page number is incorrect, Brummitt RK. 1992. Vascular plant families and genera. and it is therefore invalid (Greuter et al., 2000: Art. Surrey: Royal Botanic Gardens, Kew. 33.3). It seems that this species could also be suitably Chevalier A. 1942. Les caféiers du globe, fasc. 2: iconograhie placed in Diplospora, and it seems closely allied to des caféiers sauvages et cultivés. Encyclopédie Biologique species in the genus that have one ovule per locule 12: 1–36, pl. 1–158. (for example, D. wrayi King & Gamble; see Ali & Davis AP, Bridson DM, Jarvis C, Govaerts R. 2001. The typification and characterization of the genus Psychotria L. Robbrecht, 1991). Diplospora and Nostolachma (Rubiaceae). Botanical Journal of the Linnean Society 135: belong to a group of closely related Asian genera in 35–42. the tribe Coffeeae, together with Discospermum and Davis AP, Chester M, Maurin O, Fay M. 2007. Searching Xantonnea Pierre ex Pit. (Davis et al., 2007). It is for the relatives of Coffea (Rubiaceae, Ixoroideae): the cir- possible that all, or at least some, of these four genera cumscription and phylogeny of Coffeeae based on plastid are congeneric. Diplospora would have priority over sequence data and morphology. American Journal of Botany the other three Asian genera, and it would thus seem 94: 313–329. preferable to put Coffea viridiflora in Diplospora. Davis AP, Ruhsam M. 2005. Five new combinations and one However, Asian Coffeeae requires further critical sys- new name in Rubiaceae from South East Asia. Blumea 50: tematic investigation and a detailed taxonomic study 575–578. of the species. In this contribution, we take the oppor- Deb DB. 1988. Taxonomic revision of Mycetia Reinw. (Rubi- tunity of bringing the name N. viridiflora into current aceae) in Indian subcontinent. Bulletin of the Botanical usage only. Survey of India 28: 114–132.

Gideon O, Tjitrosoedirdjo SS, Veldkamp J-F. 1988. Notes Robbrecht E. 1999. Chassalia petitiana (Rubiaceae- on Mycetia (Rubiaceae). Floribunda 1: 17–19. Psychotrieae), an overlooked epiphytic species hidden in the Govaerts R, Frodin DG, Ruhsam M, Bridson DM, Davis African canopy. Systematic Botany 24: 315–322. AP. in prep. World checklist & bibliography of Rubiaceae. Piesschaert F, Robbrecht E, Smets E. 2001. Patterns in Surrey: Royal Botanic Gardens, Kew. pyrenes: the systematic signiﬁcance of pyrene morphology Greuter W, McNeil J, Barrie FR, Burdet HM, Demoulin in Chassalia (Rubiaceae-Psychotrieae) and related genera. V, Filgueiras TS, Nicolson DH, Silva PC, Skog JE, Flora 196: 121–131. Trehane P, Turland NJ, Hawksworth DL. 2000. Inter- Reynolds ST, Henderson RJF. 1999. Vanguerieae A. Rich. national code of botanic nomenclature (Saint Louis Code). ex Dum. (Rubiaceae) in Australia: 1. Everistia S.T. Reynolds Königstein: Koeltz Scientiﬁc Books. et R.J.F. Hend. Austrobaileya 5: 353–361.

Holmgren PK, Holmgren NH, Barnett LC. 1990. Index Robbrecht E. 1989. A remarkable new Chazaliella (African Downloaded from https://academic.oup.com/botlinnean/article/157/1/115/2418227 by guest on 23 September 2021 herbariorum in part 1: the herbaria of the world, 8th edn. Psychotrieae), exemplifying the taxonomic value of pyrene Regnum Vegetabile 120. New York: New York Botanical characters in the Rubiaceae. Bulletin Muséum National Garden. d’Histoire Naturelle, Paris, 4e Série, II, Section B, Adanso- Korthals PW. 1844. Verhandelingen over de natuurlijke nia 4: 341–349. geschiedenis der Nederlandsche overzeesche Bezittingen. Robinson CB. 1911. Botanical notes upon the land of Polillo. Botanie (Kruidkunde), Part 7. Leiden: Commissie S. en L. Philippine Journal of Science, Botany 6: 185–228. Luchtmans en C.C. van der Hoek, 197–259. Ross JH. 1975. The typification of Lycium inerme. Bothalia Lantz H, Bremer B. 2004. Phylogeny inferred from morphol- 11: 491–493. ogy and DNA data: characterizing well-supported groups in Secretariat of the Convention on Biological Diversity. Vanguerieae (Rubiaceae). Botanical Journal of the Linnean 2002. Global strategy for plant conservation. Montreal: Sec- Society 146: 257–283. retariat of the Convention on Biological Diversity. Lebrun JP, Stork AL. 1997. Énumération des plantes à Smith AC. 1988. Flora vitensis nova. A new flora of Fiji fleurs d’Afrique tropicale, Vol. 4: gamopétales, clethraceae à (spermatophytes only), Vol. 4. Kauai: Pacific Tropical Botanic lamiaceae. Geneva: Ville de Genéve editions des Conserva- Garden, Lawai. toire et Jardin Botaniques. Sohmer SH. 1988. The nonclimbing species of the genus Lobreau-Callen D, Leroy J-F. 1980. Quelques données Psychotria (Rubiaceae) in New Guinea and the Bismarck palynologiques sur le genre Coffea et autres genres du cercle Archipelago. Bishop Museum Bulletins in Botany 1: 1–339. des Caféiers. Association Scientifique Internationale du Café Sohmer SH, Davis AP. 2007. The genus Psychotria (Rubi- 9e Colloque: 479–506. aceae) in the Philippine Archipelago. SIDA, Botanical Mis- Petit E. 1964. Les espèces Africaines du genre Psychotria L. cellany 27: 1–247. (Rubiaceae) – II. Bulletin de Jardin Botanique de l’État, Turner IM. 1995. New names and new combinations in Bruxelles 36: 65–190. the Malayan flora. Asian Journal of Tropical Biology 1: Piesschaert F. 2001. Carpology and pollen morphology of the 25–27. Psychotrieae (Rubiaceae-Rubioideae). PhD Thesis. Leuven: Valeton TH. 1912. Rubiaceae. Nova Guinea 8: 767–775. Katholieke Universiteit. Verdcourt B. 1987. Notes on African Rubiaceae – Piesschaert F, Jansen S, Huysmans S, Smets E, Vanguerieae. Kew Bulletin 42: 123–199.