Nomenclatural changes and taxonomic notes
in the tribe Morindeae (Rubiaceae)
Sylvain G. RAZAFIMANDIMBISON
Birgitta BREMER
Bergius Foundation,
The Royal Swedish Academy of Sciences and Botany Department,
Stockholm University, SE-10691, Stockholm (Sweden)
sylvain.razafimandimbison@bergianska.se
birgitta.bremer@bergianska.se
Razafimandimbison S. G. & Bremer B. 2011. — Nomenclatural changes and taxonomic
notes in the tribe Morindeae (Rubiaceae). Adansonia, sér. 3, 33 (2): 283-309. DOI: 10.5252/
a2011n2a13.
KEY WORDS
Rubiaceae,
Morindeae,
Appunia,
Coelospermum,
Gynochthodes,
Morinda,
Siphonandrium,
new combinations.
ABSTRACT
A new circumscription and new generic delimitations of the tribe Morindeae
in the subfamily Rubioideae (Rubiaceae) have been proposed as a result of two
recent molecular phylogenetic studies. he adoption of a narrow circumscription of Morinda and a broad circumscription of Gynochthodes requires new
combinations and names in these genera. his study presents descriptions of the
newly delimited Gynochthodes and Morinda, and 78 new combinations (73 in
the former and five in the latter) and three new names (Gynochthodes alejandroi
Razafim. & B.Bremer, G. ridsdalei Razafim. & B.Bremer, and G. wongii Razafim. & B.Bremer). We make three lectotypifications, and recognize 15 species
in Appunia, 11 species in Coelospermum, 93 species in Gynochthodes, 39 species
in Morinda, and one species in Siphonandrium. Finally, a list of all currently
recognized species for each genus of Morindeae is presented.
ADANSONIA, sér. 3 • 2011 • 33 (2) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris.
www.adansonia.com
283
Razafimandimbison S. G. & Bremer B.
MOTS CLÉS
Rubiaceae,
Morindeae,
Appunia,
Coelospermum,
Gynochthodes,
Morinda,
Siphonandrium,
nouvelles combinaisons.
RÉSUMÉ
Changements nomenclaturaux et notes taxonomiques dans la tribu Morindeae
(Rubiaceae).
Une nouvelle circonscription et de nouvelles délimitations génériques de la tribu
des Morindeae dans la sous-famille des Rubioideae (Rubiaceae) ont été proposées
à la suite de deux récentes études phylogénétiques moléculaires. L’adoption d’une
circonscription étroite de Morinda et d’une circonscription large de Gynochthodes
nécessite de nouvelles combinaisons et de nouveaux noms dans ces genres. Cette
étude présente des descriptions de Gynochthodes et Morinda nouvellement délimités et 78 nouvelles combinations (73 chez le premier et cinq chez le second) et
trois nouveaux noms (Gynochthodes alejandroi Razafim. & B.Bremer, G. ridsdalei
Razafim. & B.Bremer et G. wongii Razafim. & B.Bremer). Nous établissons trois
lectotypifications et reconnaissons 15 espèces chez Appunia, 11 espèces chez
Coelospermum, 93 espèces chez Gynochthodes, 39 espèces chez Morinda et une
espèce chez Siphonandrium. Enfin, une liste de toutes les espèces actuellement
reconnues pour chaque genre des Morindeae est présentée.
INTRODUCTION
A new circumscription of the pantropical tribe
Morindeae of the subfamily Rubioideae (Rubiaceae)
has recently been established based on a molecular
phylogenetic study by Razafimandimbison et al.
(2008). he members of Morindeae can be diagnosed by their massive T-shaped placentae inserted
in the middle of the septum bearing two anatropous ovules per carpel and pyrenes with a single
lateral germination slit (Igersheim & Robbrecht
1993). he tribe is additionally characterized by
the frequent occurrence of head-like inflorescences
and multiple fruits. A more recent study by Razafimandimbison et al. (2009) demonstrated the
paraphyly of the type genus Morinda L. with respect
to its closely related genera Coelospermum Blume,
Gynochthodes Blume, Pogonolobus F.Muell., and
Sarcopygme Setch. & Christoph. As a consequence,
new generic limits of Morindeae were proposed in
order to make Morinda monophyletic, and that
morphologically well-defined genera were recognized in Morindeae. A total of five genera were
recognized: the Neotropical Appunia Hook.f.,
the tropical Asian and Australasian Coelospermum (including the Australian Pogonolobus), the
tropical Asian and Australasian, and Malagasy
284
Gynochthodes (including the vast majority of the
lianescent species of Morinda), the pantropical
Morinda (including Sarcopygme), and the New
Guinean Siphonandrium (see Appendix 1). Keys to
the accepted genera of Morindeae were presented
in Razafimandimbison et al. (2009). he adoption
of the new circumscriptions of Gynochthodes and
Morinda requires many new combinations. In
this paper, we present updated descriptions of the
newly circumscribed Gynochthodes and Morinda,
and make a total of 78 new combinations (73
in the former and 5 in the latter) and three new
names in Gynochthodes.
MATERIALS AND METHODS
We consulted all type specimens of the species
of Morinda at L and S herbaria, all scanned images of those at BR, BM, F, K, MO, P, and US
herbaria via the Global Plants Initiative (http://
www.bores.org) website, and those available
at the websites of A (http://www.huh.harvard.
edu) and NY (http://sciweb.nybg.org) herbaria
(all abbreviations after Holmgren et al. 1990).
In addition, we received scanned images of all
type specimens of the lianescent Morinda and
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
Sarcopygme housed at BISH herbarium. Inclusion of the species of Morinda and Sarcopygme,
respectively, in Gynochthodes and Morinda (both
sensu Razafimandimbison et al. 2009) were based
on the identification using the keys to the genera
presently recognized in Morindeae sensu Razafimandimbison et al. (2009) coupled with the results of Razafimandimbison and Bremer (unpubl.
data.) based on combined nrETS, nrITS, rps16,
and trnT-F data.
Olostyla DC., Prodromus, Systematis Naturalis 4: 440
(Sep. 1830). — Type: Olostyla corymbosa (Labill.)
DC.
SYSTEMATICS
NUMBER OF SPECIES RECOGNIZED
Eleven species (see Appendix 1).
Pogonolobus F.Muell., Fragmenta Phytographiae Australiae 1:
55 (1858). — Type: Pogonolobus reticulatus F.Muell.
Merismostigma S.Moore, Journal of the Linnean Society,
Botany 45: 332 (1921). — Type: Merismostigma neocaledonicum S.Moore.
DESCRIPTION
See Johansson (1988).
Genus Appunia Hook.f.
Genera Plantarum 2: 120 (1873). — Type: Appunia
tenuiflora (Benth.) Jack.
Bellynkxia Müll.Arg., Flora 58: 465 (1875). — Type:
Bellynkxia angulata (Benth.) Müll.Arg.
DESCRIPTION
See Burger & Taylor (1993).
NUMBER OF SPECIES RECOGNIZED
Fifteen species (see Appendix 1).
DISTRIBUTION
Neotropics.
SPECIES EXCLUDED FROM APPUNIA
Appunia parviflora Lundell., Contributions from the
University of Michigan Herbarium. Ann Arbor, MI
4: 28 (1940) = Ronabea latifolia Aubl., Histoire des
Plantes de la Guiane Française 1: 154 (1775); other
combination: Psychotria erecta (Aubl.) Standl. &
Steyerm., Publications of the Field Museum of Natural History. Botanical series 23: 24 (1943) (based
on Ronabea erecta Aubl., Histoire des Plantes de la
Guiane Française 1: 156 [1775]).
Genus Coelospermum Blume
Bijdragen tot de Flora van Nederlandsch Indië 16:
994. (Oct. 1826-Nov. 1827). — Type: Caelospermum
scandens Blume.
ADANSONIA, sér. 3 • 2011 • 33 (2)
DISTRIBUTION
Tropical Asia and Autralasia.
SPECIES EXCLUDED FROM COELOSPERMUM
1. Coelospermum barbatum Span. – Linnaea 15: 318
(1841) = Gynochthodes coriacea Blume , Bijdragen
tot de Flora van Nederlandsch Indië 16: 993 (Oct.
1826-Nov. 1827).
2. Coelospermum gmelinii Miq. – Annales Musei Lugduno-Batavi 3: 62 (1867) = Archangelica
gmelinii DC, Prodomus Systematis Naturalis IV:
170 (1830).
REMARKS
We favor the orthographic variant Coelospermum
(from coelo-, meaning with a hole in Greek) over
the original spelling Caelospermum, as the author
himself made the correction in his Flora javae
(Blume 1828). Brummitt & Taylor (1990) consider
that the spelling corrected by the original author
should be followed. he synonymy was already
established by Razafimandimbison et al. (2009:
885) for Pogonolobus.
DUBIOUS SPECIES
1. Coelospermum ahernianum Elmer. – Leaflets of
Philippine Botany 41: 3 (1906).
Remarks
We agree with Johansson (1988) that Coelospermum
ahernianum does not belong to Coelospermum,
because it does not have the characteristics of the
genus as delimited here.
285
Razafimandimbison S. G. & Bremer B.
2. Coelospermum fragrans (Montrouz.) Baill. ex Guillaumin. – Annales de l’Institut botanico-géologique
colonial de Marseille 2: 168 (1911); Figuierea fragrans Montrouz., Mémoires de l’Académie royale des
Sciences, Belles-Lettres et Arts de Lyon, Section des
Sciences 10: 220 (1860).
Remarks
Johansson (1988: 289) argued that this species
is possibly conspecific with C. monticolum Baill.
ex Guillaumin, as this latter is the only species of
Coelospermum known from the type locality of
C. fragrans (Île Art). We are unable to verify this
because we could not trace the type specimen of
C. fragrans. he location of Montrouzier material
was moved from the “Faculté de Pharmacie” to
LYJB. However, the type specimen of C. fragrans
is missing at LYJB and LY.
3. Coelospermum nigrescens (K.Krause) Guillaumin. – Archives de Botanique de Caen III (5): 41
(1929); Olostyla nigrescens K.Krause. – Botanische
Jahrbücher für Systematik, Pflanzengeschichte und
Pfanzengeographie 39: 167 (1908).
Remarks
Johnsson (1988: 291) argued that this species is possibly conspecific with either C. balansanum Baill. or
C. crassifolium J.T.Johanss. However, we were unable
to trace the type specimen of O. nigrescens, which
was presumably preserved at B herbarium.
Sciences 10: 225 (1860). — Type: Pogonanthus candollei Montrouz.
Tetralopha Hook.f., Hooker’s Icones Plantarum 11: 57, t.
1072 (1870). — Type: Tetralopha motleyi Hook.f.
Imantina Hook.f., Genera Plantarum 2: 120 (1873). —
Type: Imantina deplanchei Hook.f.
DESCRIPTION
Lianas, rarely shrubs or trees. Leaves petiolate,
chartaceous to more or less coriaceous. Inflorescences terminal, terminal on axillary (lateral)
shoots, or axillary, solitary or composed of 2 to
10 heads arranged into umbel-like, racemose or
cymose or terminal. Flowers polygamous (hermaphrodite and functionally unisexual) or dioecous, mostly completely to basally fused when
clustered in heads, sometimes in multiflorous
compound dichasia or rarely single, sessile or
sometimes pedicellate; corolla tubes shorter than
corolla lobes, rarely as long as or longer than corolla lobes; filaments inserted in corolla throat,
linear, glabrous, white, anthers partly exserted; disk
annular, persistent in fruit; styles filiform, terete;
stigmatic lobe bilobate, exserted; ovary inferior,
bilocular, with longitudinal false dissepiment
separating the ovules, ovaries of all flowers of a
head basally to completely fused; locule biovular.
Fruits fused into syncarps or simple drupes. Seeds
globose or semi-globose.
Genus Gynochthodes Blume
Bijdragen tot de Flora van Nederlandsch Indië 16: 993.
(Oct. 1826-Nov. 1827). — Type: Gynochtodes coriacea
Blume.
Guttenbergia Zoll. & Moritzi, Natuur- en Geneeskundig Archief voor Nederlandsch-Indië 2: 2
(1845), syn. nov. — Type: Guttenbergia umbellata
(L.) Zoll. & Moritzi.
Sphaerophora Blume, non (A. H. Hassall) J. Lindley
(1846), Museum Botanicum Lugduno-Batavum sive
stirpium Exoticarum 1: 179 (1850). — Type: Sphaerophora glomerata Blume. (lecto-, designated by Johansson [1994: 15], L!).
Pogonanthus Montrouz., Mémoires de l’Académie royale
des Sciences, Belles-Lettres et Arts de Lyon. Section des
286
NUMBER OF SPECIES RECOGNIZED
93 species (see Appendix 1).
DISTRIBUTION
Tropical Asia, Australasia and Madagascar.
REMARKS
We favor the orthographic variant Gynochthodes over
the original spelling Gynochtodes, as Blume himself made the correction in his Flora javae (Blume
1828). Brummitt & Taylor (1990) consider that the
spelling corrected by the original author should be
followed. Tetralopha is long established synonym of
Gynochthodes. he synonymy was already proposed
by Razafimandimbison et al. (2009: 885) for Imantina, Pogonanthus, and Sphaerophora.
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
NEW COMBINATIONS
1. Gynochthodes ammitia (Halford & A.J.Ford)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda ammitia Halford & A.J.Ford,
Austrobaileya 6: 898 (2004). — Type: Australia, Queensland, Cook Distr., Cooktown, Isabella Road, E of Isabella
Falls, c. 200 m alt., 13.III.2001, A. Ford AF 267 & J.
Holmes (holo-, BRI; iso-, DNA, NSW).
2. Gynochthodes artensis (Montrouz.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda artensis Montrouz., Mémoires
de l’Académie royale des Sciences, Belles-Lettres et Arts de
Lyon. Section des Sciences 10: 219 (1860). — Type: New
Caledonia, Île Art, Montrouzier s.n. (holo-, ?).
REMARKS
he location of Montrouzier material was moved
from the “Faculté de Pharmacie” to LYJB. However, the type specimen of M. artensis is missing at
LYJB and LY.
3. Gynochthodes badia (Y.Z.Ruan)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda badia Y.Z.Ruan, Flora Reipublicae
Popularis Sinicae 71: 327 (1999). — Type: China, Hunan,
Qianyang, 5.VI.1954, C. T. Lee 2319 (holo-, IBSC).
4. Gynochthodes bartlingii (Elmer)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda bartlingii Elmer, Leaflets of Philippine Botany 4: 1340 (1912). — Type: Philippines, Puerto
Princesa (Mt. Pulgar), Palawan, IV.1911, A. D. E. Elmer
13037 (holo-, BISH!; iso-, F!, NY!, US!).
5. Gynochthodes billardierei (Baill.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda billardierei Baill., Adansonia
n.s. 12: 231 (1879). — Type: New Caledonia, probably Balade, 1793, Labillardière s.n. (lecto-, designated
by Johansson (1994: 21), P!).
ADANSONIA, sér. 3 • 2011 • 33 (2)
6. Gynochthodes brevipes (S.Y.Hu)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda brevipes S.Y.Hu, Journal of the
Arnold Arboretum 32: 399 (1951). — Type: China,
Hainan, Po-ting, F. C. How 71754 (holo-, A!).
7. Gynochthodes bucidifolia (A.Gray)
Razafim. & B. Bremer, comb. nov.
BASIONYM. — Morinda bucidifolia A.Gray, Proceedings of the American Academy of Arts and Sciences 4: 41
(1860). — Type: Fiji, Vanua Levu, Mbua Prov., Sandalwood Bay, 1840, U.S. Expl. Exped. 62352 (holo-,
US; iso-, K, P!).
Morinda nandarivatensis Gillespie, Bernice P. Bishop
Museum Bulletin 74: 40 (1930), syn. nov. — Type:
Fiji, Viti Levu, holo North Prov., Mt. Nanggaranambuluta, E of Nandarivatu, slopes of Loma Langa
Mountain, 1200 m alt., 16.XI.1927, Gillespie 3786
(holo-, BISH!).
Morinda tripetala Christoph., Bernice P. Bishop Museum
Bulletin 154: 73 (1938), syn. nov. — Type: Samoa,
Upollu, top of Fao, open, 680 m alt., E. Christophersen
555 (holo-, BISH!, iso-, A!).
8. Gynochthodes calciphila (K.M.Wong)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda calciphila K.M.Wong, Malayan
Nature Journal 38: 97 (1984). — Type: Malaysia, Perak,
Ipok, limestone, Burkill SFN 2555 (holo-, SING).
9. Gynochthodes callicarpifolia (Y.Z.Ruan)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda callicarpaefolia Y.Z.Ruan, Flora
Reipublicae Popularis Sinicae 71: 325 (1999). —Type: China, Yunnan, Bingbian, 500 m alt., in forest, 11.VII.1953,
Mao 2517 (holo-, IBSC).
REMARKS
he specific epithet “callicarpaefolia” is changed
to “callicarpifolia” based on the Article 60.8 of
the International Code of Botanical Nomenclature
(Greuter et al. 2006).
287
Razafimandimbison S. G. & Bremer B.
10. Gynochthodes candollei (Montrouz.)
Razafim. & B.Bremer, comb. nov.
12. Gynochthodes celebica (Miq.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Pogonanthus candollei Montrouz., Mémoires
de l’Académie royale des Sciences, Belles-Lettres et Arts de
Lyon. Section des Sciences 10: 225 (1860).
BASIONYM. — Morinda celebica Miq., Flora van Nederlandsch Indië 2: 246 (1857). — Type: Celebes, among
the bushes in the coastal areas of Boni, Zollinger 3441
(holo-, L!).
Morinda candollei (Montrouz.) Beauvis., Genera Montrouzierana 57 (1901). — Type: New Caledonia, Île
Art, Montrouzier 109 (lecto-, designated by Johansson
(1994: 21), P!; isolecto-, G).
13. Gynochthodes cinnamomea (Craib)
Razafim. & B.Bremer, comb. nov.
Morinda pulchella Schltr., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pfanzengeographie 39:
267 (1906). — Type: New Caledonia, forest edges on
the mountains near Oubatche, 700 m alt., 18.XII.1902,
Schlechter 15491 (lecto-, designated by Johansson
[1994: 21], P!; isolecto-, BM, G, K).
BASIONYM. — Morinda cinnamomea Craib, Bulletin
of Miscellaneous Information, Royal Botanic Gardens,
Kew 433 (1932). — Type: Satul, Kao Kêo Range,
700 m alt., evergreen forest, Kerr 14551 (holo-, BM!;
iso-, K).
Morinda candollei Montrouz. var. subvillosa Guillaumin,
Bulletin du Muséum national d’Histoire naturelle 2:
122 (1929). — Type: New Caledonia, Prony, Franc
1853 (lecto-, designated by Johansson [1994: 23], P!;
isolecto-, A!, P!).
14. Gynochthodes cinnamomifoliata (Y.Z.Ruan)
Razafim. & B.Bremer, comb. nov.
Morinda candollei (Montrouz.) Beauvis. var. angustifolia
Guillaumin, Archives de Botanique, Mémoires 3. Caen
5: 42 (1930), syn. nov. — Type: New Caledonia, entre
Couaoua et Canala, 800 m alt., Balansa 1984 (lecto-,
designated by Johansson [1994: 23], P!).
REMARKS
We agree with Johansson (1994) that M. puchella
is a synonym of M. candollei and hereby reiterate
its synonymy in order to avoid unnecessary combinations in Gynochthodes.
11. Gynochthodes canthoides (F.Muell.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Coprosma canthoides F.Muell., Fragmenta
Phytographiae Australiae 7: 45 (1869). — Type: Australia,
South Eales, Durandoo, F. Mueller (holo-, K).
Coprosma acutifolia F.Muell. ex Benth., Flora Australiensis
3: 429 (1867), nom. illeg.
Morinda acutifolia F.Muell., Fragmenta Phytographiae
Australiae 9: 179 (1875), nom. illeg.
Morinda canthoides (F.Muell.) Halford & R.J.F.Hend.,
Austrobaileya 5: 731 (2000).
288
BASIONYM. — Morinda cinnamomifoliata Y.Z.Ruan, Flora
Reipublicae Popularis Sinicae 71: 336-337 (1999). —
Type: China, Guangxi, Guixian, Z. Z. Chen 50743
(holo-, IBSC).
15. Gynochthodes citrina (Y.Z.Ruan)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda citrina Y.Z.Ruan, Flora Reipublicae Popularis Sinicae 71: 331 (1999). — Type: China,
Guangdong, Qujiang, 3.IV.1930, S. P. Ko 50302 (holo-,
IBSC).
Morinda citrina Y.Z.Ruan var. chlorina Y.Z.Ruan, Flora
Republicae Popularis Sinicae 71: 332 (1999), syn. nov. —
Type: China, Zhejiang, Changhua, M. P. Teng 4794
(holo-, IBSC).
16. Gynochthodes cochinchinensis (DC.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda cochinchinensis DC., Prodromus
4: 449 (1830). — Type: description of Morinda umbellata
Lour., non L., Flora Cochinchinensis 140 (1790).
Morinda trichophylla Merr., Philippine Journal of Science
23: 267 (1923), syn. nov. — Type: Philippines, Yik
Tsok Mau, 17 & 19.V.1922, McClure 8755 (holo-,
A!; iso-, US!).
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
REMARKS
De Candolle’s species was based entirely on Loureiro’s
description of Morinda umbellata Lour., which is
not the same as M. umbellata L.
17. Gynochthodes collina (Schltr.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda collina Schltr., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 39: 265 (1906). — Type: New Caledonia, on
the maintain slopes near Ngoyé, 200 m alt., 30.XI.1902,
Schlechter 15154 (lecto-, designated by Johansson [1994:
33], P!; isolecto-, BM, G, K, L!, S!).
Morinda elongata Schltr., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 39:
266 (1906). — Type: New Caledonia, on the maintain
forests near Ngoyé, 400 m alt., 6.XI.1902, Schlechter
15234 (lecto-, designated by Johansson [1994: 33],
S!; isolecto-, BM!, G, K!, P!).
REMARKS
We agree with Johansson (1994) that M. elongata
is a synonym of M. collina and hereby reiterate its
synonymy in order to avoid unnecessary combinations in Gynochthodes.
18. Gynochthodes constipata
(Halford & A.J.Ford)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda constipata Halford & A.J.Ford,
Austrobaileya 81: 82 (2009). — Type: Australia, Queensland, Cook Dist., National Park Reserve 904, Wooroonooran, just S of tower 9, Mt. Bellenden Ker cableway,
17.X.2003, A. Ford AF4184 & J.Holmes (holo-, BRI;
iso-, L, K, MEL, MO, NSW, SUNIV in S!).
REMARKS
One specimen of Morinda constipata (as “Morinda
sp. 2”) was included in Razafimandimbison et al.
(2009: 881), which supported the position of this
species in Gynochthodes sensu Razafimandimbison
et al. (2009).
ADANSONIA, sér. 3 • 2011 • 33 (2)
19. Gynochthodes costata
(Merr. & L.M.Perry)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda costata Merr. & L.M.Perry,
Journal of Arnold Arboretum 26: 263 (1945). — Type:
New Guinea, 6 km southwest of Bernhard Camp,
Idenburg River, 1200 m alt., II.1939, Brass 13012
(holo-, A!).
20. Gynochthodes decipiens (Schltr.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda decipiens Schltr., Botanische
Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 39: 266 (1906). — Type: New Caledonia,
near Outbarche, 900 m alt., 17.XII.1902, Schlechter
15466 (lecto-, designated by Johansson (1994: 45), S!;
isolecto-, K, L!, P!).
21. Gynochthodes deplanchei (Hook.f.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Imantina deplanchei Hook.f., Genera
Plantarum 120 (1873); Morinda deplanchei (Hook.f.)
Baill. ex K.Schum., Die Natürlichen Pflanzenfamilien
4, 4: 138 (1891). — Type: New Caledonia, Presqu’île
de Pum, Deplanche 465 (holo-, K; iso-, P).
Morinda gatopensis Guillaumin, Archives de Botanique, Mémoires 3, Caen 5: 43 (1930). — Type: New Caledonia,
Houaïlou, XII.1869, Deplanche 407 (lecto-, designated
by Johansson [1994: 35], P!; isolecto-, A!, P!).
Morinda podocarpifolia Guillaumin, Archives de Botanique, Mémoire 3, Caen 5: 44 (1930). — Type: New
Caledonia, Houaïlou, XII.1869, Balansa 1988 (lecto-,
designated by Johansson [1994: 35], P!; isolecto-,
A!, P!).
Morinda alyxioides Guillaumin, Bulletin de la Société
botanique de France 91: 44 (1944). — Type: New Caledonia, Gatope, 1867, Vieillard s.n. (lecto-, designated
by Johansson (1994: 35), P!; isolecto-, P!).
REMARKS
We agree with Johansson (1994) that M. alyxioides,
M. gatopensis, and M. podocarpifolia are synonyms
of M. deplanchei and hereby reiterate their synonymies in order to avoid unnecessary combinations
in Gynochthodes.
289
Razafimandimbison S. G. & Bremer B.
22. Gynochthodes elliptifolia
(Quisumb. & Merr.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda elliptifolia Quisumb. & Merr.,
Philippine Journal of Science 37: 204 (1928). — Type:
Philippines, Mindanao, Davao Prov., Galintan, 250 m
alt., 24.VI.1927, Ramos & Edaño B.S. 48909 (holo-,
BR; iso-, NY!).
23. Gynochthodes elmeri (Merr.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda elmeri Merr., University of California Publications in Botany 15: 297 (1929). — Type:
Philippines, near Tawao, Elmer 21114 (holo-, UC; iso-,
F!, NY!, S!).
24. Gynochthodes gjellerupii (Valeton)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda gjellerupii Valeton, Botanische
Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 61: 152 (1927). — Type: New Guinea,
Hellwig 282 (holo-, BO).
25. Gynochthodes glaucescens (Schltr.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda glaucescens Schltr., Botanische
Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 39: 267 (1906). — Type: New Caledonia, on
the mountain slopes near Ngoyé, 300 m alt.,17.XI.1902,
Schlechter 15354 (lecto-, designated by Johansson [1994:
43], P!; isolecto-, BM, K).
Morinda schumanniana Schltr., Botanische Jahrbücher für
Systematik, Pflanzengeschichte und Pfanzengeographie
39: 268 (1906). — Type: New Caledonia, on the
mountain slopes near Ngoyé, 300 m alt., 17.XI.1902,
Schlechter 15359 (lecto-, designated by Johansson
[1994: 43], P!).
REMARKS
We agree with Johansson (1994) that M. schummaniana is a synonym of M. glaucescens and hereby
reiterate its synonymy in order to avoid unnecessary
combinations in Gynochthodes.
290
26. Gynochthodes glomerata (Blume)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Sphaerophora glomerata Blume, Museum
Botanicum Lugduno-Batavum sive stirpium Exoticarum
1: 179, f. 38 (1850); Morinda glomerata (Blume) Miq.,
Flora van Nederlandsch Indië 2: 247 (1857). — Type:
New Guinea, herb. Blume, Zippelius s.n. (lecto-, designated by Johansson [1994: 15], L!).
Sphaerophora gemella Blume, Museum Botanicum Lugduno-Batavum sive stirpium Exoticarum 1: 179, f.
36 (1850); Morinda gemella (Blume) Miq., Flora
van Nederlandsch Indië 2: 247 (1857), syn. nov. —
Type: Novâ Guineâ, herb. Blume, Zippelius s.n.
(holo-, L!).
27. Gynochthodes grayi (Seem.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda grayi Seem., Flora Vitiensis 130
(1866). — Type: Fiji, U.S. Expl. Exped., 1840, U.S.
Expl. Exped. 62353 (holo-, US; iso-, K, P).
28. Gynochthodes hainanensis
(Merr. & F.C.How)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda hainanensis Merr. & F.C.How,
Sunyatsenia 5: 188 (1940). — Type: Po-Ting Distr., TaiPing Kong, Chang-Sha Village, 330 m alt., 28.V.1935,
F. C. How 72652 (holo-, A!; iso-, NY!).
29. Gynochthodes hirtella
(Merr. & L.M.Perry)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda hirtella Merr. & L.M.Perry, Journal of Arnold Arboretum 26: 264 (1945). — Type: New
Guinea, Brass 7984 (holo-, BM!; iso-, A!, L!).
30. Gynochthodes hispida (K.M.Wong)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda hispida K.M.Wong, Malayan Nature Journal 41: 269 (1987). — Type: Malaysia Peninsula,
Johore, Padang Temambun, sandstone plateau, 600 m
alt., 27.IV.1986, Wong & Saw 34334 (holo-, KEP).
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
31. Gynochthodes hollrungiana (Valeton)
Razafim. & B.Bremer, comb. nov.
36. Gynochthodes kanalensis (Baill. ex Guillaumin)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda hollrungiana Valeton, Botanische
Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 61: 153 (1927). — Type: New Guinea,
Kaiser-Wilhems-Land, Hollrung s.n. (holo-, ?).
BASIONYM. — Morinda kanalensis Baill., Annales de
l’Institut botanico-géologique colonial de Marseille 2: 172
(1911), nom. nud.; Morinda kanalensis Baill., Bulletin
du Muséum national d’Histoire naturelle, sér. 2, 1: 118
(1929), nom. nud.; Morinda kanalensis Baill. ex Guillaumin, Archives de Botanique, Mémoires 3, Caen 5:
43 (1930). — Type: New Caledonia, Mt. Humboldt,
12.X.1869, Balansa 1987 (lecto-, designated by Johansson (1994: 53), P!; isolecto-, P!).
REMARKS
We have not been able to trace the original material.
According to Stafleu & Cowan (1986: 649), the
type specimen may be located at BO. We made a
request to this herbarium without success.
32. Gynochthodes howiana (S.Y.Hu)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda howiana S.Y.Hu, Journal of Arnold Arboretum 32: 400 (1951). — Type: China, Hainan,
Poting, 14.IV.1935, F. C. How 71911 (lecto-, designated
by Liu & Zhang [2008: 578], A!).
33. Gynochthodes hupehensis (S.Y.Hu)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda hupehensis S.Y.Hu, Journal of
Arnold Arboretum 32: 400 (1951). — Type: China,
Hupeh, En-shih, H. C. Chow 1818 (holo-, A!).
34. Gynochthodes jackiana (Korth.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda jackiana Korth., Nederlandsch
Kruidkundig Archief 2, 2: 227 (1851). — Type: Indonesia,
Borneo, Korthals s.n. (holo-, L; iso-, A!).
35. Gynochthodes jasminoides (A.Cunn.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda jasminoides A.Cunn., Botanical
Magazine 61: t. 3351 (1834). — Type: Australia, Colony
of Port Jackson, Cunningham s.n. (holo-, K).
ADANSONIA, sér. 3 • 2011 • 33 (2)
37. Gynochthodes lacunosa
(King & Gamble)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda lacunosa King & Gamble, Journal
of the Asiatic Society of Bengal 73: 87 (1904). — Types:
Perak, King’s collector 4320, King’s collector 6030, King’s
collector 8254 (Syn-, CAL).
38. Gynochthodes leparensis (Valeton)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda leparensis Valeton, Icones Bogorienses 4: t. 362 (1913). — Type: Lepar, Teysmann,
Hortus Bogor, cult. XVII C 102 (holo-, L!).
39. Gynochthodes leptocalama (Wernham)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda leptocalama Wernham, Journal
of Botany, British & Foreign. London 56: 77 (1918). —
Type: New Guinea, Mt. Gawada, 4000 ft., Forbes 864
(holo-, BM!; iso-, K, L!).
40. Gynochthodes litseifolia (Y.Z.Ruan)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda litseifolia Y.Z.Ruan., Flora Reipublicae Popularis Sinicae 71: 335 (1999). — Type: China,
Jiangxi, Jiangang-mountain, 1300 m alt., 09.VII.1965,
S. S. Lai 3488 (holo-, IBSC).
291
Razafimandimbison S. G. & Bremer B.
41. Gynochthodes micrantha (Valeton)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda micrantha Valeton, Botanische
Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 61: 153 (1927). — Type: New Guinea,
800 m alt., VIII.1910, L. Schultze 43 (holo-, ?).
REMARKS
We have not been able to trace the original material.
According to Stafleu & Cowan (1986: 649), the
type specimen may be located at BO. We made a
request to this herbarium without success.
42. Gynochthodes microcephala (Bartl. ex DC.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda microcephala Bartl. ex DC.,
Prodromus 4: 449 (1830). — Type: Phillipines, Luzon
Island, T. Haenke s.n. (holo-, G-DC!).
43. Gynochthodes mollis (A.Gray)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda mollis A.Gray, Proceedings of the
American Academy of Arts and Sciences 4: 41 (1860). —
Type: Fiji, U.S. Expl. Exped. 62377 (holo-, US; iso-,
K, NY!).
Morinda candollei Montrouz. var. villosa Guillaumin,
Archives de Botanique, Mémoires 3, Caen 5: 42 (1930),
syn. nov. — Type: New Caledonia, Gatope, Vieillard 2741 pro parte (lecto-, designated by Johansson
[1994: 27], P!).
44. Gynochthodes montana (J.T.Johanss.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda montana J.T.Johanss., Opera
Botanica 122: 49 (1994). — Type: New Caledonia,
Mont Panié, 1400 m alt., 28.XI.1983, McPherson 6090
(holo-, MO; iso-, S!).
45. Gynochthodes myrtifolia (A.Gray)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda myrtifolia A.Gray, Proceedings of the American Academy of Arts and Sciences 4:
292
41 (1860); Morinda myrtifolia A.Gray var. myrtifolia
(autonym used by J.T.Johanss., Opera Botanica 122: 55
[1994]). — Type: Fiji, U.S. Expl. Exped. 62351 (holo-,
US; iso-, K, NY!).
Morinda forsteri Seem., Flora Vitiensis 129 (1866). —
Type: Society Islands, J. R. & G. Forster s.n. (lecto-,
designated by Smith & Darwin [1988: 333], K; isolecto-, BM).
Morinda choriophylla Baill., Adansonia s.n. 12: 246
(1879); Morinda myrtifolia A.Gray var. choriophylla (Baill.) J.T.Johnss., Opera Botanica 122: 57
(1994). — Type: New Caledonia, à l’embouchure
de la rivière du Houaïlou, XII.1869, Balansa 1989
(lecto-, designated by Johansson [1994: 57], P!;
isolecto-, K, P!).
Morinda glandulosa Merr., Philippine Journal of Science.
Section C, Botany 9: 146 (1914); Morinda umbellata
L. var. glandulosa Merr., Occasional Papers of Bernice
Pauahi Bishop Museum of Polynesian Ethnology and
Natural History 15: 220 (1940). — Type: Marianas
Islands, Tumon Road, XI.1911, Guam Experiment
Station 37, J. B. hompson 376 (lecto-, designated by
Johansson (1994: 55), NY!).
Morinda velutina Guillaumin, Archives de Botanique,
Mémoires 3, Caen 5: 44 (1930); Morinda myrtifolia
A.Gray var. velutina (Guillaumin) J.T.Johanss., Opera Botanica 122: 59 (1994), syn. nov. —Type: New
Caledonia, above Ouroué, near the mouth of Dotio,
I.1872, Balansa 3421 (lecto-, designated by Johansson
(1994: 59), P!; isolecto-, A!, P!).
Morinda forsteri Seem. subsp. guillauminii Däniker,
Vierteljahrsschrift der Naturforschenden Gesellschaft
in Zürich 78: 465 (1933). — Type: New Caledonia,
groves on the Nouméa peninsula, X.1870-I.1871,
Balansa 2992 (holo-, Z; iso-, P!).
Morinda myrtifolia A.Gray var. brevifolia J.T.Johanss., Opera Botanica 122: 55 (1994). —Type: New Caledonia,
Négropo, northern plateau of Mont Prokoméo, 700 m
alt., 27.I.1972, MacKee 24947 (holo-, P!).
REMARKS
We agree with Johansson (1994) that M. choriophylla, M. glandulosa, and M. forsteri are synonyms
of M. myrtifolia and hereby reiterate their synonymies in order to avoid unnecessary combinations
in Gynochthodes.
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
46. Gynochthodes nanlingensis (Y.Z.Ruan)
Razafim. & B.Bremer, comb. nov.
50. Gynochthodes officinalis (F.C.How)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda nanlingensis Y.Z.Ruan, Flora
Reipublicae Popularis Sinicae 71: 329 (1999). — Type:
China, Hunan, Yizhang, 5.VI.1941, Q. Liu 2948 (holo-,
IBSC).
BASIONYM. — Morinda officinalis F.C.How, Acta Phytotaxonomica Sinica 7: 326-327 (1958). — Type: China,
Kwangtung, Lofou Shan, 11.V.1930, N. K. Chun 40965
(holo-, IBSC).
Morinda nanlingensis Y.Z.Ruan var. pauciflora Y.Z.Ruan,
Flora Reipublicae Popularis Sinicae 71: 331 (1999),
syn. nov. — Type: China, Zhejiang, Hanzhou, X. Y.
He 20528 (holo-, IBSC).
Morinda officinalis F.C.How var. hirsuta F.C.How, Acta
Phytotaxonomica Sinica 7: 328 (1958), syn. nov. —
Type: China, Hainan, Tungfang Hsien, 15.XII.1955,
Hainan Exped. 287 (holo-, IBSC).
Morinda nanlingensis Y.Z.Ruan var. pilophora Y.Z.Ruan,
Flora Reipublicae Popularis Sinicae 71: 331 (1999),
syn. nov. — Type: China, Hunan, Huitong, Z. T. Li
2887 (holo-, IBSC).
47. Gynochthodes neocaledonica (S.Moore)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Lucinaea neocaledonica S.Moore, Journal
of the Linnean Society, Botany 45: 328 (1921); Morinda
neocaledonica (S.Moore) Guillaumin, Archives de Botanique, Mémoire 3, Caen 3: 44 (1930). — Type: New
Caledonia, Taom, 500 m alt., 2.XII.1914, Compton
2304 (lecto-, designated by Johansson (1994: 27),
BM; iso-, P!).
48. Gynochthodes nitida (Merr.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda nitida Merr., Philippine Journal
of Science 17: 435 (1920, publ. 1921). — Type: Philippines, Bucas Grande Island, VI.1919, Ramos & Pascasio
BS 35074 (holo-, A!; iso-, US!).
49. Gynochthodes oblongifolia (Valeton)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda oblongifolia Valeton, Botanische
Jahrbücher für Systematik, Pflanzengeschichte und Pfanzengeographie 61: 153 (1927). — Type: New Guinea, 100 m
alt., 22.V.1912, Ledermann 7366 (holo-, ?).
REMARKS
We have not been able to trace the original material.
According to Stafleu & Cowan (1986: 649), the
type specimen may be located at BO. We made a
request to this herbarium without success.
ADANSONIA, sér. 3 • 2011 • 33 (2)
51. Gynochthodes oligocephala
(Merr. & L.M.Perry)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda oligocephala Merr. & L.M.Perry,
Journal of Arnold Arboretum 26: 265 (1945). — Type:
New Guinea, Tarar, Wassi Kussa River, XII.1936, Brass
8490 (holo-, A; iso-, BM!, L!, NY!).
52. Gynochthodes parvifolia (Bartl. ex DC.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda parvifolia Bartl. ex DC., Prodromus
4: 449 (1830). — Type: Philippines, Manila, 1.XII.1912,
T. Haenke s.n. (holo- PR or PRC?; iso- MO!).
Morinda cumingiana S.Vidal, Phanerogamae Cumingianae
Philippinarum: 184 (1885); Lucinaea cumingiana
S.Vidal, Phanerogamae Cumingianae Philippinarum:
216 (1885), syn. nov. — Type: Philippines, Albay
Prov., H. Cuming 1242 (holo-, BM; iso-, L).
REMARKS
he holotype of Gynochthodes parvifolia is not found
at G-DC herbarium or G herbarium. De Candolle
(1830) referred to the Haenke herbarium, where he
himself saw the type specimen. So, it is possible that
the holotype is at PR or PRC herbarium, where the
collections of Haenke are housed. We made requests
from these herbaria without success.
53. Gynochthodes phyllireoides (Labill.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda phyllireoides Labill., Sertum austrocaledonicum 49: t. 49 (1825). — Type: New Caledonia,
293
Razafimandimbison S. G. & Bremer B.
probably Balade, 1793, Labillardière s.n. (lecto-, designated
by Johansson (1994: 37), P!; iso-, BM, G, G-DC, K).
57. Gynochthodes pubiofficinalis (Y.Z.Ruan)
Razafim. & B.Bremer, comb. nov.
Morinda vieillardii Baill., Adansonia s.n. 12: 230 (1879). —
Type: New Caledonia, Wagap, 1861, Vieillard 719 pro
parte (lecto-, designated by Johansson (1994: 37), P!;
isolecto-; P!).
BASIONYM. — Morinda pubiofficinalis Y.Z.Ruan, Flora
Reipublicae Popularis Sinicae 71: 334 (1999). — Type:
China, Guizhou, Tongren, Tsiang 5990 (holo-, IBSC).
Morinda fallax Schltr., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 39: 267
(1906). — Type: New Caledonia, in the mountain
forests near Yaouhé, 400 m alt., 15.X.1902, Schlechter
15034 (lecto-, designated by Johansson (1994: 37),
P!; isolecto-, BM, G, K, L!, P!, Z).
Morinda ligustrina S.Moore, Journal of the Linnean Society, Botany 14: 335 (1921). — Type: New Caledonia,
Ignambi, 1400 m alt., 01.VIII.1914, Compton 1580
(holo-, BM).
REMARKS
We agree with Johansson (1994) that M. fallax, M. ligustrina, and M. vieillardi are synonyms of M. phyllireoides and hereby reiterate their synonymies in order to
avoid unnecessary combinations in Gynochthodes.
54. Gynochthodes platyphylla (Merr.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda platyphylla Merr., Philippine
Journal of Science 11: 33 (1916). — Type: Sorsogon
Prov., Luzon Island, VIII.1915, M. Ramos BS 23414
(holo-, A!; iso-, BISH!, L!, NY!; US!).
55. Gynochthodes podistra (Halford & A.J.Ford)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda podistra Halford & A.J.Ford, Austrobaileya 6: 895 (2004). — Type: Australia, Queenland,
Cook District, Daintree River National Park, Black Mountain area, Daintree River headwaters, 25.V.1998, P. I. Forster
PIF22959, R. Jago, R. Jensen & R. Booth (holo-, BRI).
56. Gynochthodes polyneura (Miq.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda polyneura Miq., Flora van Nederlandsch Indië 2: 244 (1857). — Types: Horsfield s.n.
(syn-, BM, K, U in L), Junghuhn s.n. (syn-, L, U in L).
294
58. Gynochthodes reticulata (Valeton)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Lucinaea reticulata Valeton, Contribution to the
Phytogeography and Flora of the Arfak Mountains 180 (1917);
Morinda reticulata (Valeton) Valeton, Botanische Jahrbücher
für Systematik, Pflanzengeschichte und Pfanzengeographie 61:
152 (1927). — Type: New Guinea, Irian Jaya, Arfak Mts,
Angi lakes, 7000 ft, Gibbs 5580 (holo-, BM!).
59. Gynochthodes retropila (Halford & A.J.Ford)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda retropila Halford & A.J.Ford, Austrobaileya 81: 85 (2009). — Type: Australia, Wooroonooran
National Park, start of Bartle Frere walking track, 27.XI.2001,
P.I.Forster PIF27757 & A.M.Young (holo-, BRI).
REMARKS
One specimen of Morinda retropila (as Morinda sp. 1)
was included in Razafimandimbison et al. (2009),
which supported the position of this species in Gynochthodes sensu Razafimandimbison et al. (2009).
60. Gynochthodes retusa (Poir.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda retusa Poir., Encyclopédie méthodique, Botanique Supplément 4: 316 (1797). — Type:
Madagascar, unknown s.n. (holo-, P-LA).
61. Gynochthodes ridleyi (King & Gamble)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda umbellata L. var. ridleyi King &
Gamble, Journal of the Asiatic Society of Bengal, Part 2,
Natural History 73: 88 (1904); Morinda ridleyi (King &
Gamble) Ridl., Flora of the Malay Peninsula 2: 118
(1923). — Types: Singapore, in Botanic Garden Jungle, Ridley 5668, Ridley 6470, Ridley 6471, and Ridley
6916 (Syn-, K).
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
62. Gynochthodes rigida (Miq.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda rigida Miq., Flora van Nederlandsch
Indië 2: 246 (1857). — Type: Indonesia, Sumatra, near
Sibogo, Teysmann s.n. (lecto-, designated by Johansson
(1994: 15), U in L).
63. Gynochthodes rugulosa (Y.Z.Ruan)
B.Bremer, comb. nov.
BASIONYM. — Morinda rugulosa Y.Z.Ruan, Flora Reipublicae Popularis Sinicae 71: 328 (1999). — Type: China,
Guangxi, Gongcheng, 13.VII.1931, S. S. Sin. 128651
(holo-, IBSC).
64. Gynochthodes sarmentosa (Blume)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda sarmentosa Blume, Bijdragen tot
de Flora van Nederlandsch Indië 1006 (Oct. 1826-Nov.
1827). — Type: Indonesia, Java, Blume s.n. (holo-,
L!).
Morinda sarmentosa Blume var. puberula Miq., Flora van
Nederlandsch Indië 2: 245 (1857), syn. nov. — Type:
Indonesia, Java, Horsfield s.n. (holo-, L or U in L?).
65. Gynochthodes scabrifolia (Y.Z.Ruan)
Razafim. & B.Bremer, comb. nov.
67. Gynochthodes subcaudata (Valeton)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda subcaudata Valeton, Botanische
Jahrbücher für Systematik, Pflanzengeschichte und Pfanzengeographie 61: 154 (1927). — Types: Ledermann 9069,
9885, 9925 (syn-, ?).
REMARKS
We have not been able to trace the original material.
According to Stafleu & Cowan (1986: 649), the
type specimens may be located at BO. We made a
request to this herbarium without success.
68. Gynochthodes triandra (S.Moore)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda triandra S.Moore, Journal of
Botany 61 (Supplement): 25 (1923). — Type: Sogere,
Forbes 487 (holo-, BM).
69. Gynochthodes trimera (Hillebr.)
Razafim & B.Bremer, comb nov.
BASIONYM. — Morinda trimera Hillebr., Flora Hawaiian
Islands 177 (1888). — Type: Hawaiian Islands, east Maui,
Hamakua, J. M. Lydgate s.n. (holo-, BISH!).
Morinda sandwicensis O.Deg., Flora Hawaiiensis 332
(1936); Morinda sandwicensis O.Deg. var. sandwicensis, (autonym used by H.St.John, Pacific Science
33 (4): 372 [1979, publ. 1980]), syn. nov. — Type:
Hawaiian Island, Waianae Mts. 10.VII.1932, O. Degener, K. Park, & Y. Nitta 9423 (holo-, BISH!; iso-,
MO!, NY!).
BASIONYM. — Morinda scabrifolia Y.Z.Ruan, Flora Reipublicae Popularis Sinicae 71: 332 (1999). — Type: China,
Yunnan, Bingbian, China-Soviet Union Exped. 3515
(holo-, IBSC).
Morinda lainaiensis H.St.John, Pacific Science 33 (4):
369 (1979, publ. 1980), syn. nov. — Type: Hawaiian
Islands, Lanai Island, Kahinahina, 14.VI.1915, G. C.
Mubro 262 (holo-, BISH!).
66. Gynochthodes shuanghuaensis
(C.Y.Chen & M.S.Huang)
Razafim. & B.Bremer, comb. nov.
Morinda sandwicensis O.Deg. var. glabrata (O.Deg.) H.St.
John, Pacific Science 33 (4): 372 (1979, publ. 1980);
Morinda sandwicensis O.Deg. f. glabrata O.Deg., Flora
Hawaiiensis, Family 332 (1936), syn. nov. — Type:
Hawaiian Islands, Waianae Mts., 13.V.1932, O. Degener, K. Park, D. Topping, & O. Swezey 4150 (holo-,
BISH!; iso-, NY).
BASIONYM. — Morinda shuanghuaensis C.Y.Chen &
M.S.Huang, Acta Phytotaxonomica Sinica 14: 70-71
(1976). — Type: China, Guangdong, Wuhua, 25.X.1975,
Baji Expedition 045 (holo-, IBSC).
ADANSONIA, sér. 3 • 2011 • 33 (2)
Morinda sandwicensis O.Deg. var. hosakae H.St.John, Pacific Science 33 (4): 374 (1979, publ. 1980), syn. nov. —
Type: Hawaiian Islands, Oahu Island, Koolau Range,
295
Razafimandimbison S. G. & Bremer B.
16.IV.1923, E. Y. Hosaka 963 (holo-, BISH!; isotype,
NTBG!).
Morinda waikapuensis H.St.John, Pacific Science 33 (4):
376 (1979, publ. 1980), syn. nov. — Type: Hawaiian
Islands, Maui Island, Waikapu Valey, VI.1910, C. N.
Forbes 133.M (holo-, BISH!).
REMARKS
Morinda trimera is one of the two arborescent species of Morinda that we transferred to Gynochthodes
sensu Razafimandimbison et al. (2009) based on
Razafimandimbison & Bremer (unpubl. data)
analyses based on combined nrETS, nrITS, rps16,
and trnT-F data. he range of its flowers length
also falls within that of Gynochthodes. We consider
M. lainaiensis, M. sandwicensis, and M. waikapuensis
to be synonyms of M. trimera and hereby provide
their synonymies in order to avoid unnecessary
combinations in Gynochthodes.
70. Gynochthodes truncata (J.T. Johanss.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda truncata J.T.Johanss., Opera
Botanica 122: 41 (1994). — Type: New Caledonia, sommet nord du Mt Kaala, 1000-1050 m alt., 25.XII.1966,
MacKee 16146 (holo-, P!).
71. Gynochthodes umbellata (L.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda umbellata L., Species Plantarum
176 (1753). — Type: Herb. Hermann 3: 11, no. 81 (BM000621833) (lecto-, designated by Smith & Darwin
[1988: 333], BM).
Morinda padavara Juss. ex Schult., Systema Vegetabilium
5: 216 (1819), syn. nov. — Type: Malabar s.n. (lecto-,
designated here, P-JU!).
Morinda scandens Roxb., Flora of India 2: 201 (1824),
syn. nov. — Type: drawing nº 2570 in Icones Roxburghinae (holo-, K).
Morinda umbellata L. var. scortechinii King & Gamble,
Journal of the Asiatic Society of Bengal, Part 2. Natural History 73: 88 (1904); Morinda scortechinii (King & Gamble) Ridl., Flora of the Malay Peninsula 2: 118 (1923),
syn. nov. — Type: Scortechini 2015 (holo-, K).
296
Morinda umbellata L. var. tonkinensis Pit., in Lecomte
H. & Humbert H., Flore Générale de l’ Indochine 3:
323 (1924), syn. nov. — Type: Vietnam, Lang-bian,
A. Chevalier s.n. (lecto-, designated here, P!).
Morinda umbellata L. var. accuminatissima Valeton,
Botanische Jahrbücher für Systematik, Pflanzengeschichte
und Pfanzengeographie 61: 153 (1927), syn. nov. —
Type: New Guinea, 850 m atl., Ledermann 9331
(holo-, ?).
Morinda boninensis Ohwi, Feddes Repertorium 36: 57
(1934); Morinda umbellata L. var. boninensis (Ohwi)
T.Yamaz., Journal of Japanese Botany 61: 343 (1986),
syn. nov. — Type: S. Nishimura s.n. (holo-, TO).
Morinda umbellata L. var. archboldiana Fosberg, Sargentia 1: 123 (1942), syn. nov. — Type: Fiji, Viti Levu,
Serua, vicinity of Ngaloa, hulanuku, dense forest,
30 m alt., 29.IV.1941, Degener 15115 (holo-, US!;
iso-, A!, NY!, S!).
Morinda umbellata L. var. forsteri Seem., Sargentia 1: 123
(1942), syn. nov. — Type: Society Islands, J. R. & G.
Forsteri s.n. (lecto-, designated by Smith & Darwin
(1988: 333), K!; isolecto-, BM!).
Morinda umbellata L. var. hahazimensis T.Yamaz., Journal
of Japanese Botany 61: 343 (1986), syn. nov. — Type:
Japan, Hahazima Island, 15.VII.1920, T. Nakai s.n.
(holo-, TI).
Morinda umbellata L. var. obovata Y.Z.Ruan, Flora
Reipublicae Popularis Sinicae 71: 325 (1999), syn.
nov. — Type: China, Guangdong, Fengshun & G. Li
201473 (holo-, IBSC).
REMARKS
We have not been able to trace the original material of Morinda umbellata L. var. accuminatissima
Valeton. According to Stafleu and Cowan (1986:
649), the type specimen may be located at BO, P,
and U. We made requests to these herbaria without success.
72. Gynochthodes villosa (Hook.f.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda villosa Hook.f., Flora of British India 3: 158 (1880). — Type: Wallich’ Cat. 8425
(holo-, K-W).
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
73. Gynochthodes wallichii (Kurz)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Morinda wallichii Kurz, Journal of the
Asiatic Society of Bengal, Part 2, Natural History 41: 313
(1872). — Type: Herb. Kurz, Tenasserim s.n. (holo-,
CAL).
NEW NAMES
1. Gynochthodes alejandroi
Razafim. & B.Bremer, nom. nov.
REPLACED NAME. — Morinda philippinensis Elmer,
Leaflets of Philippine Botany 3: 1044 (1911). — Type:
Philippines, Capiz Prov., Magallanes (Mt. Giting-giting),
Sibuan Island, V.1910, A. D. E. Elmer 12460 (holo-, A!;
iso-, L!, NY!; US!).
REMARKS
he specific epithet “philippinensis” cannot be used
in combination with Gynochthodes, as it would be
a later homonym of Gynochthodes philippinensis
(Elmer) Merr. he replacement name alejandroi
refers to Dr. Grecebio J. Alejandro, who has a good
knowledge of the Philippian Rubiaceae.
2. Gynochthodes ridsdalei
Razafim. & B.Bremer, nom. nov.
REPLACED NAME. — Morinda reticulata Gamble, Bulletin
of Miscellaneous Information, Royal Botanic Gardens, Kew
248 (1920). — Type: India (south), evergreen forests of
Travancore, at Merchiston, 600 m alt., IV.1895, Bourdillon 591 (lecto-, K, designated here).
REMARKS
Morinda reticulata Gamble is an illegitimate name, as
it is a later homonym of Morinda reticulata Benth.,
which has priority over the former. Razafimandimbison et al. (2009) confirmed that the Australian
Morinda reticulata Benth. belongs to Coelospermum,
consistent with Baillon (1879) who already made
the new combination Coelospermum decipiens Baill.
he specific epithet “reticulata” cannot be used in
combination with Gynochthodes, as it would be a
later homonym of Gynochthodes reticulata (Valeton)
Razafim. & B.Bremer. he replacement name rids-
ADANSONIA, sér. 3 • 2011 • 33 (2)
dalei refers to Dr. Colin Ridsdale, who has a great
knowledge of the tropical Asian Rubiaceae. Gamble
(1920) listed two specimens, Bourdillon 591 and
M. Rama Row 1281, when describing his Morinda
reticulata. We chose Bourdillon 591 as lectotype.
3. Gynochthodes wongii
Razafim. & B.Bremer, nom. nov.
REPLACED NAME. — Morinda coriacea Merr., Philippine
Journal of Science 17: 435 (1920, publ. 1921). — Type:
Mindanao (Surigao), Ramos and Pascasio B.S. 34439
(holo-, A!; US!).
REMARKS
he specific epithet “coriacea” cannot be used in
combination with Gynochthodes, as it would be a
later homonym of Gynochthodes coriacea Blume, the
type of Gynochthodes. he replacement name wongii
refers to Professor Khoon Meng Wong (Malaysia),
who has a great knowledge of the Southeast Asian
Rubiaceae.
Genus Morinda L.
Species plantarum: 176 (1753). — Type: Morinda royoc
L. (lecto-, designated by Hitchcock: 132 [1929]).
Rojoc Adans., Famille des Plantes 2: 246 (1763).
Appunettia Good, Journal of Botany 64 (Supplement 2):
30 (1926). — Type: Appunettia angolensis Good.
Sarcopygme Setch. & Christoph., Occasional Papers of
Bernice Pauahi Bishop Museum of Polynesian Ethnology
and Natural History 11: 4 (1935), syn. nov. — Type:
Sarcopygme pacifica (Reinecke) Setch. & Christoph.
DESCRIPTION
Small trees or shrubs, rarely lianas. Leaves petiolate
(rarely subsessile), chartaceous to coriaceous, glabrous. Inflorescences mostly leaf-opposed (modified terminal inflorescences), sometimes axillary
with heads arranged into secondarily compound
inflorescences, composed of 1 to 3 heads, sometimes axillary with heads arranged into secondarily compound inflorescences, pedunculate (rarely
sessile), branched. Flowers bisexual; corolla tubes
297
Razafimandimbison S. G. & Bremer B.
much larger than corolla lobes; filaments inserted in
corolla throat, linear, glabrous, white; anthers partly
exserted (rarely well exserted); disk annular, persistent in fruit; style filiform, terete; stigma bilobate,
more or less recurved, not exserted; ovaries inferior,
bilocular, with longitudinal false dissepiment, ovaries of all flowers of a head basally to completely
fused; locule biovular. Fruits fused into syncarps.
Seed globose or semi-globose.
NUMBER OF SPECIES RECOGNIZED
40 species (see Appendix 1).
DISTRIBUTION
3. Morinda multinervis (Setch. & Christoph.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Sarcopygme multinervis Setch. & Christoph.,
Bernice P. Bishop Museum Bulletin 154: 69 (1938). — Type:
Samoa, Christophersen & Hume 2299 (holo-, BISH!).
4. Morinda pacifica (Reinecke)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Sarcocephalus pacificus Reinecke, Botanische Jahrbücher für Systematik, Pflanzengeschichte und
Pfanzengeographie 25: 681, pl. 13 (1898); Sarcopygme
pacifica (Reinecke) Setch. & Christoph., Occasional
Papers of Bernice Pauahi Bishop Museum of Polynesian
Ethnology and Natural History 11: 4 (1935). — Type:
Samoa, Upolu, Reinecke 1623 (holo-, HBG).
Pantropical.
NEW COMBINATIONS
he type of the Samoan genus Sarcopygme was nested
in Morinda sensu stricto in Razafimandimbison
et al. (2009). As a consequence, we transfer below
all the five described species to Morinda.
1. Morinda intermedia (Setch. & Christoph.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Sarcopygme intermedia Setch. & Christoph.,
Bernice P. Bishop Museum Bulletin 154: 67 (1938). —
Type: Samoa, wet forest above Matavanu crater, atl.
1400 m, 28.VII.1931 (fr.), Christophersen & Hume 2193
(holo-, BISH!).
5. Morinda ramosa (Lauterb.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Sarcocephalus ramosus Lauterb., Botanische Jahrbücher für Systematik, Pflanzengeschichte und
Pfanzengeographie 41: 235 (1908); Sarcopygme ramosa
(Lauterb.) Setch. & Christoph., Occasional Papers of Bernice Pauahi Bishop Museum of Polynesian Ethnology and
Natural History 11: 4 (1935). — Type: Samoa, Savaii,
Mataana, alt. 1600 m, 20.IX.1906, Vaupel 525 (holo-,
B probably destroyed).
SPECIES EXCLUDED FROM MORINDA
A total of 29 species of Morinda have been excluded from Morinda sensu Razafimandimbison
et al. (2009), of which 12 neotropical species are
transferred back to Appunia.
1. Morinda axillaris Lam. ex Poir.
2. Morinda mayorii (Setch.)
Razafim. & B.Bremer, comb. nov.
BASIONYM. — Breonia (?) mayorii Setch., American Samoa:
44, pls 6-7 (1924); Sarcopygme mayorii (Setch.) Setch. &
Christoph., Occasional Papers of Bernice Pauahi Bishop
Museum of Polynesian Ethnology and Natural History 11:
5 (1935). — Type: Samoa, Tutuila, trail to Matafao,
I.1922, Setchell 419 (holo-, UC).
298
Encyclopédie Méthodique. Botanique Supplément 4:
315 (1797) = Saldinia axillaris (Lam. ex Poir.) Bremek.,
Candollea 16: 110 (1957).
2. Morinda aurantiaca (K.Krause) Steyerm.
Memoirs of the New York Botanical Garden 23: 385
(1972) = Appunia aurantiaca (K.Krause) Sandwith,
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
Kew Bulletin 472 (1931) (based on Rudgea aurantiaca
K.Krause, Notizblatt des Botanichen Gartens und Museums
zu Berlin-Dahlem 6: 206 [1914]).
3. Morinda beccariana Baill. ex K.Schum.
Die Natürlichen Pflanzenfamilien 4, 4: 138 (1891) =
Primatomeris beccariana (Baill. ex K.Schum.) J.T.Johanss.,
Opera Botanica 94: 57 (1987).
4. Morinda borneensis (Baill.) K.Schum.
Die Natürlichen Pflanzenfamilien 4, 4: 138 (1891) =
Rennellia borneensis Baill., Bulletin mensuel de la Société
linnéenne de Paris 1: 205 (1879).
5. Morinda brachycalyx (Bremek.) Steyerm.
Memoirs of the New York Botanical Garden 23: 385
(1972) = Appunia brachycalyx (Bremek.) Steyerm., Memoirs
of the New York Botanical Garden 17: 359 (1967) (based
on Bellynkxia brachycalyx Bremek., Recueil des Travaux
botaniques néerlandais 31: 227 [1934]).
9. Morinda guatemalensis
(Donn.Sm.) Steyerm.
Memoirs of the New York Botanical Garden 23: 385
(1972) = Appunia guatemalensis Donn.Sm., Botanical
Gazette 48: 294 (1909).
10. Morinda hypotephra F.Muell.
Victorian Naturalist 6: 55 (July 1889) = Palmeria
hypotephra (F.Muell.) Domin (Monimiaceae), Repertorium Specierum Novarum Regni Vegetabilis 12: 390
(1913).
REMARKS
he name Morinda hypotephra F.Muell. was misapplied in the past (up to 1997) to the species
that we refer to an undescribed Morinda sp. 1 in
Razafimandimbison et al. (2009) and was recently
described by Halford & Ford (2009) as Morinda
retropila Halford & A.J.Ford (see Appendix 1).
he type of the name Morinda hypotephra F.Muell.
is actually a species of Palmeria, P. hypotephra
(F.Muell.) Domin, of the family Monimiaceae
and this synonymy was confirmed by Whiffin &
Foreman (2007).
6. Morinda calycina (Benth.) Steyerm.
Memoirs of the New York Botanical Garden 23: 385
(1972) = Appunia calycina (Benth.) Sandwith, Kew
Bulletin 471 (1931) (based on Coffea calycina Benth.,
Journal of Botany by William Jackson Hooker 3: 232
[1841]; other combination: Bellynkxia calycina (Benth.)
Bremek., Recueil des Travaux botaniques néerlandais 31:
276 [1934]).
11. Morinda korthalsiana Miq.
Annales Musei Botanici Lugduno-Batavi 4: 214 (1869)
= Rennellia morindiformis (Korth.) Ridl., Bulletin of
Miscellaneous Information, Royal Botanic Gardens, Kew
1939: 609 (1940).
7. Morinda debilis (Sandwith) Steyerm.
12. Morinda longiloba
(Steyerm.) Steyerm.
Memoirs of the New York Botanical Garden 23: 385
(1972) = Appunia debilis Sandwith, Kew Bulletin 471
(1931).
Memoirs of the New York Botanical Garden 23: 385
(1972) = Appunia longiloba Steyerm., Memoirs of the
New York Botanical Garden 17: 359 (1967).
8. Morinda esquirolii H.Lév.
13. Morinda longipedunculata Steyerm.
Flore du Kouy-Tchéou 368 (1915) = Macaranga andamanica Kurz, Forest Flora of British Burma 2: 389
(1877).
Annals of Missouri Botanical Garden 75: 340 (1988) =
Appunia longipedunculata (Steyerm.) Delprete, Journal of
Botanical Research Institute of Texas 3: 809 (2009).
ADANSONIA, sér. 3 • 2011 • 33 (2)
299
Razafimandimbison S. G. & Bremer B.
14. Morinda muscosa Jacq.
21. Morinda sparsiflora (Steyerm.) Steyerm.
Selectarum Stirpium Americanarum Historia in qua ad
Linnaeanum Systema Determinatae 65 (1763) = Psychotria muscosa (Jacq.) Steyerm., Memoirs of the New York
Botanical Garden 23: 671 (1972).
Memoirs of the New York Botanical Garden 23: 385
(1972) = Appunia sparsiflora Steyerm., Memoirs of the
New York Botanical Garden 17: 356 (1967).
22. Morinda speciosa Wall. ex Kurz
15. Morinda odontocalyx (Sandwith) Steyerm.
Memoirs of the New York Botanical Garden 23: 385
(1972) = Appunia odontocalyx Sandwith, Kew Bulletin
472 (1931) (based on Bellynksia odontocalyx (Sandwith)
Bemek., Recueil des Travaux botaniques néerlandais 31:
276 [1934]).
16. Morinda palmetorum DC.
Prodromus 4: 448 (1830) = Psychotria peduncularis
var. palmetorum (DC.) Verdc., Kew Bulletin 30: 257
(1975).
17. Morinda peduncularis Kunth
Nova Genera et Species Plantarum 3: 380 (1818) = Appunia peduncularis (Kunth) Delprete, Journal of Botanical
Research Institute of Texas 3: 809 (2009).
Forest Flora of Burma 2: 62 (1877) = Rennellia speciosa (Wall. ex Kurz) Hook.f., Flora of British India 3:
158 (1880).
23. Morinda sumatrana Miq.
Annales Musei Botanici Lugduno-Batavi 4: 213 (1869) =
Rennellia elliptica Korth., Nederlandsch Kruidkundig
Archief 2, 2: 257 (1851).
24. Morinda surinamensis (Bremek.) Steyerm.
Memoirs of the New York Botanical Garden 23: 386
(1972) = Appunia surinamensis (Bremek.) Steyerm.,
Memoirs of the New York Botanical Garden 17: 359 (1967)
(based on Bellynkxia surinamensis Bremek., Recueil des
Travaux botaniques néerlandais 31: 276 (1934); other
combination: Cordiera surinamensis Miq. ex Bremek.,
Recueil des Travaux botaniques néerlandais 31: 276 [1934]
nom. inval. pro. syn.).
18. Morinda polysperma Jack
Malayan Miscellanies 1: 14 (1820) = Schradera polysperma (Jack) Puff, Buchner & Greimler, Blumea 43: 293
(1998); other combination: Lucinaea polysperma (Jack)
K.Schum., Die Natürlichen Pflanzenfamilien 4, 4: 64
[1891].
19. Morinda salomoniensis Engl.
Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pfanzengeographie 7: 478 (1886) = Coelospermum salomoniense (Engl.) J.T.Johanss., Blumea
33: 281 (1988).
20. Morinda seibertii (Standl.) Steyerm.
Memoirs of the New York Botanical Garden 23: 386
(1972) = Appunia seibertii Standl., Annals Missouri Botanical Garden 24: 208 (1937).
300
25. Morinda tenuiflora (Benth.) Steyerm.
Memoirs of the New York Botanical Garden 23: 386
(1972) = Appunia tenuiflora (Benth.) B.D.Jacks., Kew Index
1: 166 (1893) (based on Coffea tenuiflora Benth., Journal
of Botany by William Jackson Hooker 3: 232 (1841)).
26. Morinda tribrachya K.Schum.
Die Naturlichen Pflanzenfamilien 4(4): 138 (1891) =
Rennellia morindiformis (Korth.) Ridl., Bulletin of Miscellaneous Information, Royal Botanic Gardens, Kew 1939:
609 (1940).
27. Morinda triphylla (Ducke) Steyerm.
Memoirs of the New York Botanical Garden 23: 386
(1972) = Appunia triphylla Ducke, Notizblatt des Bota-
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
nischen Gartens und Museums zu Berlin-Dahlem 11: 482
(1932; other combination: Bellynkxia triphylla (Ducke)
Bremek., Recueil des Travaux botaniques néerlandais 31:
276 (1934)).
28. Morinda volubilis Merr.
Philippine Journal of Science, Section C, Botany 1 (Supplement 5): 137 (1906) = Coelospermum volubile (Merr.)
J.T.Johanss., Blumea 33: 281 (1988).
29. Morinda venezuelensis (Steyerm.) Steyerm.
Memoirs of the New York Botanical Garden 23: 386
(1972) = Appunia venezuelensis Steyerm. Memoirs of the
New York Botanical Garden 17: 356 (1967).
DUBIOUS SPECIES
1. Morinda lastelliana Baill. – Adansonia 12: 231
(1879).
Remarks
Examination of the type specimen of M. lastelliana
(at P) shows that it probably represents a species of
the genus rubiaceous Danais in the tribe Danaideae
(A. P. Davis, pers. com.).
Genus Siphonandrium K.Schum.
Nachträge zur Flora der deutschen Schutzgebiete in der
Südsee 394 (1905). — Type: Siphonandrium intricatum
K.Schum.
DESCRIPTION
See Schumann & Lauterbach (1905).
NUMBER OF RECOGNIZED SPECIES
One species (see Appendix 1).
DISTRIBUTION
New Guinea.
Acknowledgements
We thank David Halford for informing us about the
identity of Morinda hypotephra; Sachiko Nishida and
Armand Randrianasolo for providing some Japanese
ADANSONIA, sér. 3 • 2011 • 33 (2)
and Chinese literature; Martin Callmander for checking the presence of the type specimens of Morinda
cochinchinensis DC and Morinda parvifolia Bartl. ex
DC at G-DC and G herbaria; Martin Callmander and
Nicolas Fumeaux for helping with the nomenclatural issues of M. cochinchinensis; Aaron Davis, Elmar
Robbrecht, and Rafaël Govaerts for their invaluable
comments on an early version of the paper; and the
following herbaria for making scanned images of their
type specimens of Morinda available online: A, BR,
BM, F, K, L, MO, NY, P, U in L, US; S and UPS
for allowing access to collections; and Dr. Neill Snow
and BISH for providing scanned images.
REFERENCES
AUBLET J. B. C. F. 1775. — Histoire des plantes de la
Guiane Française 1. P.-F. Didot jeune, London & Paris,
392 p.
BAILLON, H. E. 1879. — Sur l’Imantina. Bulletin mensuel
de la Société linnéene de Paris 1: 202-205.
BLUME C. L. 1826-1827. — Rubiaceae. Bijdragen tot de
Flora van Nederlandsch Indië 16: 913-1018.
BLUME C. L. 1828. — Enumeratio plantarum Javae et
Insularum adjacentium minus cognitarum vel novarum.
Fasciculus II. Lugduni Batavorum, Leiden, v-viii.
BREMEKAMP, C. E. B. 1934. — Notes on the Rubiaceae
of Surinam. Recueil des Travaux botaniques néerlandais
31: 248-308.
BREMEKAMP C. E. B. 1957. — Monographie du genre
Saldinia A. Rich. (Rubiacées). Candollea 16: 91-129.
BRUMMITT R. K. & TAYLOR N. P. 1990. — To correct or
not to correct? Taxon 39: 298-306.
BURGER W. & TAYLOR C. M. 1993. — Family # 202
Rubiaceae, in BURGER W. (ed.), Flora Costaricensis.
Fieldiana Botany 33: 1-333.
CANDOLLE A. DE 1830. — Prodomus Systematis Naturalis
IV. Treuttel et Würtz, Paris, 683 p.
DELPRETE P. G. 2009. — Rubiaceae, in FEUILLET C. (ed.),
Checklist of the plants of the Guianas Shield – 1. Update
to the Angiosperms. Journal of the Botanical Research
Institute of Texas 3: 809-813.
DOMIN K. 1913. — Eighth contribution to the Flora of
Australia. Repertorium Specierum Novarum Regni Vegetabilis 12: 388-390.
DUCKE A. 1932. — Neue arten aus der Hylaea Brasiliens.
Notizblatt des Botanischen Gartens und Museums zu
Berlin-Dahlem 11: 471-483.
HALFORD D. A. & FORD A. J. 2009. — Two new species
of Morinda L. (Rubiaceae) from north-east Queensland.
Austrobaileya 8: 81-90.
HIERN W. P. 1898. — Catalogue of the African Plants col-
301
Razafimandimbison S. G. & Bremer B.
lected by Dr. Friedrich Welwitsch in 1853-61, Part 3.
Longman and Co, London: 511-784.
HOLMGREN P. K., HOLMGREN N. H. & BARNETT L. C.
1990. — Index Herbariorum. New York Botanical
Garden, New York, 693 p.
HOOKER W. J. 1841. — Coffea calycina. Journal of Botany,
British and Foreign 3: 232.
HOOKER J. D. 1880. — he Flora of British India 3L. Reeve
and Co., London, 610 p.
HITCHCOCK A. S. 1929. — Proposal by Hichcock A.
S. (Washington) and Green M. L. (Kew) - Standardspecies of Linnean genera of Phanerogamae (1753-54),
in RAMSBOTTOM J., SPRAGUE T. A., WILLMOTT A. J. &
WAKEFIELD E. M. (eds), International Botanical Congress,
Cambridge, 1930, Nomenclature: Proposal by British
Botanist. Wyman & Sons, London: 110-199.
JACKSON B. D. 1893. — Appunia tenuiflora (Benth.)
B.D.Jacks. Index Kewensis 1: 166.
JOHANSSON J. T. 1987. — Revision of the genus Prismatomeris hw. (Rubiaceae, Morindeae). Opera Botanica 94: 1-62.
JOHANSSON J. T. 1988. — Revision of the genus Caelospermum Blume (Rubiaceae, Rubioideae, Morindeae).
Blumea 33: 265-297.
JOHANSSON J. T. 1994. — he genus Morinda (Morindeae,
Rubioideae, Rubiaceae) in New Caledonia: Taxonomy
and phylogeny. Opera Botanica 122: 5-67.
KORTHALS P. W. 1851. — Overzigt der Rubiaceën van
de Nederlandschoostindische Kolonien. Nederlandsch
Kruidkundig Archief 2: 255-257.
KRAUSE K. 1908. — Olostyla nigrescens K.Krause. Botanische Jahrbücher für Systematik, Pflanzengeschichte und
Pfanzengeographie 40: 39.
KRAUSE K. 1914. — Rubiaceae. Notizblatt des Botanichen
Gartens und Museums zu Berlin-Dahlem 6: 200-212.
KUTZ W. S. 1877. — Forest flora of British Burma 2. Office of the Superintendent of Government Printing,
Calcutta, 1162 p.
LÉVEILLÉ A. A. H. 1914-1915. — Flore du Kouy-Tchéou.
Le Mans, 532 p.
MCNEILL J., BARRIE F. R., BURDET H. M., DEMOULIN V.,
HAWKSWORTH D. L., MARHOLD K., NICOLSON D. H.,
SILVA P. C., SKOG J. E., WIERSEMA J. H. & TURLAND N.
J. 2006. — International Code of Botanical Nomenclature 2006 (Vienna Code): adopted by the Seventeenth
International Botanical Congress, Vienna, Austria, July
2005. Regnum Vegetabile 146. 1-568.
MIQUEL F. A. G. 1867. — Coelospermum gmelinii Miq.
Annals Musei Botanici Lugduno-Batavi 3: 62.
PUFF C., GREIMLER J. & BUCHNER R. 1998. — Revision
of Schradera (Rubiaceae-Schradereae) in Malesia. Blumea 43: 287-335.
RAZAFIMANDIMBISON S. G., MCDOWELL T. D., HALFORD
D. A. & BREMER B. 2009. — Molecular phylogenetics
and genetic assessment in the tribe Morindeae (RubiaceaeRubioideae): How to circumscribe Morinda L. to be
302
monophyletic? Molecular Phylogenetics and Evolution
52: 879-886.
RAZAFIMANDIMBISON S. G., RYDIN C. & BREMER B.
2008. — Evolution and trends in the Psychotrieae alliance
(Rubiaceae): a rarely reported evolutionary change of
many-seeded carpels from one-seeded carpels. Molecular
Phylogenetics and Evolution 48: 207-223.
RIDLEY N. H. 1930. — Notes on some Malayan Rubiaceae.
Morindeae. Bulletin of Miscellaneous Information, Royal
Botanic Gardens, Kew 9: 600-611.
SANDWITH N. Y. 1931. — Contribution to the Flora of
tropical America: VIII. Bulletin of Miscellaneous Information, Royal Botanic Gardens, Kew 10: 467-492.
SCHUMANN K. 1891. — Rubiaceae, in ENGLER A. &
PRANTL K. (eds). Die natürlichen Pflanzenfamilien vol.
IV(4). W. Engelmann, Leipzig: 1-156.
SCHUMANN K. & LAUTERBACH K. 1905. — Nachträge zur
Flora der Deutschen Schutzgebiete in der Südsee. Gebrüder
Borntraeger, Leipzig, 446 p.
SETCHELL W. A. & CHRISTOPHERSEN E. 1935. — Preliminary notes on Sarcopygme, a new rubiaceous genus
from Samoa. Occasional Papers of Bernice Pauahi Bishop
Museum of Polynesian Ethnology and Natural History
11: 3–5.
SPANOGHE J. B. 1841. — Prodromus florae timorensis. Linnaea 15: 318.
STAFLEU F. A. & COWAN R. S. 1986. — Taxonomic literature. A selective guide to botanical publications and
collections with dates, commentaries and types. 2nd ed.,
6 (Sti-Vuy). Regnum Vegetabile 115: 1-926.
STANDLEY P. C. & STEYERMARK J. A. 1943. — Study of
Central America Plants. III. Field Museum of Natural
History, Botanical Series 23: 24.
STEYERMARK J. A. 1967. — Rubiaceae, in MAGUIRE B.
& COLLABORATORS (eds), he botany of the Guayana
Highland. Memoirs of the New York Botanical Garden
17: 1-439.
STEYERMARK, J. A. 1972. — Rubiaceae, in MAGUIRE B.
& COLLABORATORS (eds), he botany of the Guayana
Highland – Part IX. Memoirs of the New York Botanical
Garden 23: 227–832.
VALETON, T. 1927. — Die Rubiaceae von Papuasien 2.
Coffeoideae, in LAUTERBACH K. (ed.), Beiträge zur Flora
von Papuasien 14. Botanische Jahrbücher für Systematik,
Pflanzengeschichte und Planzengeographie 61: 32-163.
VERDCOURT B. 1975. — Studies in the Rubiaceae-Rubioideae for the flora of tropical East Africa. Kew Bulletin
30: 247-326.
WHIFFIN T. & FOREMAN D. B. 2007. — Monimiaceae.
Flora of Australia 2. Australian Biological Resources
Study/CSIRO Publishing, 486 p.
Submitted on 29 October 2009;
accepted on 17 February 2011.
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
APPENDIX 1
Index of names associated with Appunia, Coelospermum, Gynochthodes, Morinda, and Siphonandrium. 1, new names; bold, type
species.
Species recognized in this study
Homotypic synonyms
Heterotypic synonyms
Appunia angulata (Benth.) Baill.
Ixora angulata Benth.; Bellynkxia
angulata (Benth.) Müll.Arg.
Appunia aurantiaca (K.Krause)
Rudgea aurantiaca K.Krause; Morinda
Sandwith
aurantiaca (K.Krause) Steyerm.
Appunia brachycalyx (Bremek.)
Bellynkxia brachycalyx Bremek.;
Steyerm.
Morinda brachycalyx (Bremek.)
Steyerm.
Appunia calycina (Benth) Sandwith
Coffea calycina Benth.; Bellynkxia
calycina (Benth.) Bremek.; Morinda
calycina (Benth.) Steyerm.
Appunia debilis Sandwith
Morinda debilis (Sandwith) Steyerm.
Appunia guatemalensis Donn.Sm.
Morinda guatemalensis (Donn.Sm.)
Steyerm.
Appunia longipedunculata (Steyerm.) Morinda longipedunculata Steyerm.
Delprete
Appunia megalantha C.M.Taylor &
Lorence
Appunia odontocalyx Sandwith
Bellynkxia odontocalyx (Sandwith)
Bremek.; Morinda odontocalyx
(Sandwith) Steyerm.
Appunia peduncularis (Kunth)
Morinda peduncularis Kunth
Delprete
Appunia siebertii Standl.
Morinda siebertii (Standl.) Steyerm.
Appunia surinamensis (Bremek.)
Bellynkxia surinamensis Bremek.;
Steyerm.
Morinda surinamensis (Bremek.)
Steyerm.; Cordiera surinamensis Miq.
ex Bremek.
Coffea tenuiflora Benth.; Morinda
Appunia longiloba Steyerm.; Morinda
Appunia tenuiflora (Benth.)
tenuiflora (Benth.) Steyerm.
longiloba (Steyerm.)Steyerm.;
B.D.Jacks.
Appunia sparsiflora Steyerm.; Morinda
sparsiflora (Steyerm.)Steyerm.
Appunia triphylla Ducke
Bellynkxia triphylla (Ducke) Bremek.;
Morinda triphylla (Ducke) Steyerm.
Appunia venezuelensis Steyerm.
Morinda venezuelensis (Steyerm.)
Steyerm.
Coelospermum balansanum Baill.
Stylocorina corymbosa Labill.;
Olostyla corymbosa (Labill.) DC.;
C. corymbosum (Labill.) Baill. ex
K.Schum.; C. billardieri Däniker;
Merismostigma neocaledonicum
Moore
Coelospermum crassifolium
J.T.Johanss.
Coelospermum dasylobium Halford &
A.J.Ford
Coelospermum decipiens Baill.
Morinda reticulata Benth.
Coelospermum monticolum Baill. ex
Guillaumin
Coelospermum paniculatum F.Muell.
Coelospermum purpureum Halford &
A.J.Ford
ADANSONIA, sér. 3 • 2011 • 33 (2)
303
Razafimandimbison S. G. & Bremer B.
APPENDIX 1 – Continuation.
Species recognized in this study
Coelospermum reticulatus (F.Muell.)
Benth.
Coelospermum salomoniense (Engl.)
J.T.Johanss.
Coelospermum truncatum (Wall.)
Baill. ex K.Schum.
Coelospermum volubile (Merr.)
J.T.Johanss.
Gynochthodes alejandroi Razafim. &
B.Bremer1
Gynochthodes ammitia (Halford &
A.J.Ford) Razafim. & B.Bremer
Gynochthodes artensis (Montrouz.)
Razafim. & B.Bremer
Gynochthodes australensis
J.T.Johanss.
Gynochthodes badia (Y.Z.Ruan)
Razafim. & B.Bremer
Gynochthodes bartlingii (Elmer)
Razafim. & B.Bremer
Gynochthodes billardierei (Baill.)
Razafim. & B.Bremer
Gynochthodes brevipes (S.Y.Hu.)
Razafim. & B.Bremer
Gynochthodes bucidifolia (A.Gray)
Razafim. & B.Bremer
Homotypic synonyms
Pogonolobus reticulatus F.Muell.
Heterotypic synonyms
Morinda salomoniensis Engl.
Morinda volubilis auct. non Merrill
Cupia truncata (Wall.) DC.; Diplospora Coelospermum scandens Blume;
truncata (Wall.) Steud.; Pseudixora
C. corymbosum Blume ex DC.;
truncata (Wall.) Miq.; Trisciadia
C. morindiforme Pierre ex Pitard;
truncata (Wall.) Hook.f.
C. acuminatum Geddes; C. luteum
Geddes; C. kanehirae Merr.
Morinda volubile Merr.
Morinda philippinensis Elmer
Morinda ammitia Halford & A.J.Ford
Morinda artensis Montrouz.
Morinda badia Y.Z.Ruan
Morinda bartlingii Elmer
Morinda billardierei Baill.
Morinda brevipes S.Y.Hu
Morinda bucidifolia A.Gray; Morinda
umbellata L. var. bucidifolia (A.Gray)
Forsberg
Gynochthodes calciphila (K.M.Wong) Morinda calciphila K.M.Wong
Razafim. & B.Bremer
Gynochthodes callicarpifolia
Morinda callicarpifolia Y.Z.Ruan
(Y.Z.Ruan) Razafim. & B.Bremer
Gynochthodes candollei (Montrouz.) Pogonanthus candollei Montrouz.;
Razafim. & B.Bremer
Morinda candollei (Montrouz.)
Beauvis.
Gynochthodes canthoides (F.Muell.)
Razafim. & B.Bremer
Gynochthodes celebica (Miq.)
Razafim. & B.Bremer
Gynochthodes cinnamomea (Craib)
Razafim. & B.Bremer
Gynochthodes cinnamomifoliata
(Y.Z.Ruan) Razafim. & B.Bremer
Gynochthodes citrina (Y.Z.Ruan)
Razafim. & B.Bremer
Gynochthodes cochinchinensis (DC.)
Razafim. & B.Bremer
304
Morinda pulchella Schltr.; Morinda
candollei Montrouz. var. subvillosa
Guillaumin; Morinda candollei
(Montrouz.) Beauvis. var. angustifolia
Guillaumin
Coprosma canthoides F.Muell.;
Coprosma acutifolia F.Muell. ex
Benth.; Morinda acutifolia F.Muell.;
Morinda canthoides (F.Muell.)
Halford & R.J.F.Hend
Morinda celebica Miq.
Morinda cinnamomea Craib
Morinda cinnamomifoliata Y.Z.Ruan
Morinda citrina Y.Z.Ruan
Morinda cochinchinensis DC.
Morinda citrina Y.Z.Ruan var. chlorina
Y.Z.Ruan
Morinda trichophylla Merr.
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
APPENDIX 1 – Continuation.
Species recognized in this study
Gynochthodes collina (Schltr.)
Razafim. & B.Bremer
Gynochthodes constipata (Halford &
A.J.Ford) Razafim. & B.Bremer
Gynochthodes coriacea Blume
Gynochthodes costata (Merr. &
L.M.Perry) Razafim. & B.Bremer
Gynochthodes decipiens (Schltr.)
Razafim. & B.Bremer
Gynochthodes deplanchei (Hook.f.)
Razafim. & B.Bremer
Homotypic synonyms
Morinda collina Schltr.
Heterotypic synonyms
Morinda elongata Schltr.
Morinda constipata Halford & A.J.Ford
Morinda costata Merr. & L.M.Perry
Morinda decipiens Schltr.
Imantina deplanchei Hook.f.; Morinda Morinda gatopensis Guillaumin;
deplanchei (Hook.f.) Baill.
Morinda podocarpifolia Guillaumin;
Morinda alyxioides Guillaumin
Gynochthodes elliptifolia (Quisumb. & Morinda elliptifolia Quisumb. & Merr.
Merr.) Razafim. & B.Bremer
Gynochthodes elmeri (Merr.)
Morinda elmeri Merr.
Razafim. & B.Bremer
Gynochthodes epiphytica (Rech.)
Plectronia epiphytica Rech.
Gynochthodes ovalifolia (Valeton)
A.C.Sm. & S.P.Darwin
Kaneh.; G. trukensis Hosok.
Gynochthodes gjellerupii (Valeton)
Morinda gjellerupii Valeton
Razafim. & B.Bremer
Gynochthodes glaucescens (Schltr.) Morinda glaucescens Schltr.
Morinda schumanniana Schltr.
Razafim. & B.Bremer
Gynochthodes glomerata (Blume)
Sphaerophora glomerata Blume;
Sphaerophora gemella Blume;
Razafim. & B.Bremer
Morinda glomerata (Blume) Miq.
Morinda gemella (Blume) Miq.
Gynochthodes grayi (Seem.)
Morinda grayi Seem.
Razafim. & B.Bremer
Gynochthodes hainanensis (Merr. & Morinda hainanensis Merr. & F.C.How
F.C.How) Razafim. & B.Bremer
Gynochthodes hirtella (Merr. &
Morinda hirtella Merr. & L.M.Perry
L.M.Perry) Razafim. & B.Bremer
Gynochthodes hispida (K.M.Wong)
Morinda hispida K.M.Wong
Razafim. & B.Bremer
Gynochthodes hollrungiana (Valeton) Morinda hollrungiana Valeton
Razafim. & B.Bremer
Gynochthodes howiana (S.Y.Hu)
Morinda howiana S.Y.Hu
Razafim. & B.Bremer
Gynochthodes hupehensis (S.Y.Hu) Morinda hupehensis S.Y.Hu
Razafim. & B.Bremer
Gynochthodes jackiana (Korth.)
Morinda jackiana Korth.
Razafim. & B.Bremer
Gynochthodes jasminoides (A.Cunn.) Morinda jasminoides A.Cunn.
Razafim. & B.Bremer
Gynochthodes kanalensis (Baill. ex
Morinda kanalensis Baill. ex
Guillaumin) Razafim. & B.Bremer
Guillaumin
Gynochthodes lacunosa (King &
Morinda lacunosa King & Gamble
Gamble) Razafim. & B.Bremer
Gynochthodes lenticellata (C.B.Rob.) Tetralopha lenticellata C.B.Rob.
Merr.
Gynochthodes leparensis (Valeton)
Morinda leparensis Valeton
Razafim. & B.Bremer
Gynochthodes leptocalama
Morinda leptocalama Wernham
(Wernham) Razafim. & B.Bremer
Gynochthodes litseifolia (Y.Z.Ruan)
Morinda litseifolia Y.Z.Ruan.
Razafim. & B.Bremer
ADANSONIA, sér. 3 • 2011 • 33 (2)
305
Razafimandimbison S. G. & Bremer B.
APPENDIX 1 – Continuation.
Species recognized in this study
Homotypic synonyms
Gynochthodes macrophylla Kurz
Gynochthodes micrantha (Valeton)
Morinda micrantha Valeton
Razafim. & B.Bremer
Gynochthodes microcephala (Bartl. Morinda microcephala Bartl. ex DC.
ex DC.) Razafim. & B.Bremer
Gynochthodes mindanaensis Merr.
Gynochthodes mollis (A.Gray)
Morinda mollis A.Gray
Razafim. & B.Bremer
Gynochthodes montana (J.T.Johanss.) Morinda montana J.T.Johanss.
Razafim. & B.Bremer
Gynochthodes myrtifolia (A.Gray)
Morinda myrtifolia A.Gray; M.
Razafim. & B.Bremer
myrtifolia A.Gray var. myrtifolia
J.T.Johanss.
Gynochthodes nanlingensis
(Y.Z.Ruan) Razafim. & B.Bremer
Morinda nanlingensis Y.Z.Ruan
Gynochthodes neocaledonica
(S.Moore) Razafim. & B.Bremer
Lucinaea neocaledonica S.Moore;
Morinda neocaledonica (S.Moore)
Guillaumin
Tetralopha nigra Merr.
Morinda nitida Merr.
Gynochthodes nigra (Merr.) Merr.
Gynochthodes nitida (Merr.)
Razafim. & B.Bremer
Gynochthodes oblongifolia (Valeton)
Razafim. & B.Bremer
Gynochthodes officinalis (F.C.How)
Razafim. & B.Bremer
Gynochthodes oligantha Merr.
Gynochthodes oligocephala (Merr. &
L.M.Perry) Razafim. & B.Bremer
Gynochthodes oresbia Halford &
A.J.Ford
Gynochthodes parvifolia (Bartl. ex
DC.) Razafim. & B.Bremer
Gynochthodes phylippinensis (Elmer)
Merr.
Gynochthodes phyllireoides (Labill.)
Razafim. & B.Bremer
Gynochthodes platyphylla (Merr.)
Razafim. & B.Bremer
Gynochthodes podistra (Halford &
A.J.Ford) Razafim. & B.Bremer
Gynochthodes pollilensis (C.B.Rob.)
Ruhsam
Gynochthodes polyneura (Miq.)
Razafim. & B.Bremer
Gynochthodes proboscidea Pierre
ex. Pit.
Gynochthodes puberula Craib
Gynochthodes pubifolia Merr.
306
Heterotypic synonyms
Morinda candollei Montrouz. var.
villosa Guillaumin
Morinda forsteri Seem.;
M. choriophylla Baill.; M. myrtifolia
A.Gray var. choriophylla (Baill.)
J.T.Johanss.; M. glandulosa Merr.;
M. velutina Guillaumin; M. forsteri
Seem. subsp. guillauminii Däniker;
M. myrtifolia A.Gray var. brevifolia
J.T.Johanss.
Morinda nanlingensis Y.Z.Ruan var.
pauciflora Y.Z.Ruan; M. nanlingensis
Y.Z.Ruan var. pilophora Y.Z.Ruan
Morinda oblongifolia Valeton
Morinda officinalis F.C.How
Morinda officinalis var. hirsuta
F.C.How
Morinda oligocephala Merr. &
L.M.Perry
Morinda parvifolia Bartl. ex DC.
Morinda cumingiana S.Vidal
Tetralopha philippinensis Elmer
Morinda phyllireoides Labill.
Morinda vieillardii Baill.; Morinda fallax
Schltr.; Morinda ligustrina S.Moore
Morinda platyphylla Merr.
Morinda podistra Halford & A.J.Ford
Tetralopha pollilensis C.B.Rob.
Morinda polyneura Miq.
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
APPENDIX 1 – Continuation.
Species recognized in this study
Homotypic synonyms
Gynochthodes pubiofficinalis
Morinda pubiofficinalis Y.Z.Ruan
(Y.Z.Ruan) Razafim. & B.Bremer
Gynochthodes reticulata (Valeton)
Lucinaea reticulata Valeton; Morinda
Razafim. & B.Bremer
reticulata (Valeton) Valeton
Gynochthodes retropila (Halford &
Morinda retropila Halford & A.J.Ford
A.J.Ford) Razafim. & B.Bremer
Gynochthodes retusa (Poir.)
Morinda retusa Poir.
Razafim. & B.Bremer
Gynochthodes ridleyi (King & Gamble) Morinda umbellata L. var. ridleyi
Razafim. & B.Bremer
King & Gamble;
Morinda ridleyi king & Gamble
Gynochthodes ridsdalei Razafim. &
Morinda reticulata Gamble
B.Bremer1
Gynochthodes rigida (Miq.)
Morinda rigida Miq.
Razafim. & B.Bremer
Gynochthodes rugulosa (Y.Z.Ruan)
Morinda rugulosa Y.Z.Ruan
B.Bremer
Gynochthodes sarmentosa (Blume)
Morinda sarmentosa Blume
Razafim. & B.Bremer
Gynochthodes scabrifolia (Y.Z.Ruan) Morinda scabrifolia Y.Z.Ruan
Razafim. & B.Bremer
Gynochthodes sessilis Halford
Gynochthodes shuanghuaensis
Morinda shuanghuaensis C.Y.Chen &
(C.Y.Chen & M.S.Huang) Razafim. & M.S.Huang
B.Bremer
Gynochthodes subcaudata (Valeton) Morinda subcaudata Valeton
Razafim. & B.Bremer
Gynochthodes sublanceolata Miq.
Gynochthodes triandra (S.Moore)
Morinda triandra S.Moore
Razafim. & B.Bremer
Gynochthodes trimera (Hillebr.)
Morinda trimera Hillebr.
Razafim & B.Bremer
Gynochthodes truncata (J.T. Johanss.) Morinda truncata J.T.Johanss.
Razafim. & B.Bremer
Gynochthodes umbellata (L.)
Morinda umbellata L.
Razafim. & B.Bremer
Heterotypic synonyms
Morinda sarmentosa Blume var.
puberula Miq.
Gynochthodes tetrandra Kuntze
Morinda sandwicensis O.Deg.;
M. lainaiensis H.St.John;
M. sandwicensis O.Deg. var. glabrata
O.Deg.; M. sandwicensis O.Deg. var.
hosakae H.St.John; M. sandwicensis
O.Deg. var. sandwicensis;
M. waikapuensis H.St.John
Morinda padavara Juss. ex Schult.;
M. scandens Roxb.; M. umbellata
L. var. scortechinii King & Gamble;
M. umbellata L. var. tonkinensis Pit.;
M. umbellata L. var. accuminatissima
Valeton; M. boninensis Ohwi;
M. umbellata L. var. archboldiana
Fosberg; M. umbellata L. var.
bucidifolia (A.Gray) Forsberg;
M. umbellata L. var. forsteri Seem.;
M. umbellata L. var. hahazimensis
T.Yamaz.; M. umbellata L. var.
obovata Y.Z.Ruan
Gynochthodes verticillata (Valeton)
Hosok
ADANSONIA, sér. 3 • 2011 • 33 (2)
307
Razafimandimbison S. G. & Bremer B.
APPENDIX 1 – Continuation.
Species recognized in this study
Gynochthodes villosa (Hook.f.)
Razafim. & B.Bremer
Gynochthodes wallichii (Kurz)
Razafim. & B.Bremer
Gynochthodes wongii Razafim. &
B.Bremer1
Morinda angolensis (R.D.Good)
F.White
Morinda angustifolia Roxb.
Morinda asteroscepa K.Schum.
Morinda buchii Urb.
Morinda citrifolia L.
Homotypic synonyms
Morinda villosa Hook.f.
Morinda wallichii Kurz
Morinda coriacea Merr.
Appunettia angolensis R.D.Good
Morinda squarrosa Buch.-Ham.
Morinda aspera Wight & Am.;
M. asperula Standl.; M. bracteata
(Roxb.) Kurz; M. chachuca
Buch.-Ham.; M. ligulata Blanco;
M. littoralis Blanco; M. macrophylla
Desf.; M. mudia Buch.-Ham.;
M. multiflora Roxb.; M. nodosa
Buch.-Ham.; M. quadrangularis
G.Don; M. stenophylla Spreng.;
M. teysmanniana Noronha; M. tinctoria
Noronha; M. zollingeriana Miq.
Morinda exserta Roxb.; M. tinctoria
Roxb.
Morinda coreia Buch.-Ham.
Morinda corneri K.M.Wong
Morinda elliptica (Hook.f.) Ridl.
Morinda citrifolia L. var. elliptica
Hook.f.
Morinda fasciculata Benth.
Morinda geminata DC.
Morinda hoffmannioides Standl.
Morinda intermedia (Setch. &
Christoph.) Razafim. & B.Bremer
Morinda latibracteata Valeton
Morinda laxa DC.
Morinda leiantha Kurz
Morinda longiflora G.Don
Morinda longifolia Craib
Morinda longissima Y.Z.Ruan
Morinda lucida Benth.
Morinda mayorii (Setch.) Razafim. &
B.Bremer
Morinda moaensis Alain
Morinda morindoides (Baker) MilneRedh.
Morinda multinervis (Setch. &
Christoph.) Razafim. & B.Bremer
Morinda nana Craib.
Morinda pacifica (Reinecke)
Razafim. & B.Bremer
Heterotypic synonyms
Morinda batesii Wernham,
M. chrysorhiza (Thonn.) DC.
Sarcopygme intermedia Setch. &
Christoph.
Morinda citrifolia sensu Hiern (1898)
Breonia (?) mayorii Setch.;
Sarcopygme mayorii (Setch.) Setch. &
Christoph.
Morinda confusa Huch.
Sarcopygme multinervis Setch. &
Christoph.
Sarcocephalus pacificus Reinecke;
Sarcopygme pacifica (Reinecke)
Setch. & Christoph.
Morinda panamensis Seem.
Morinda pandurifolia Kuntze
308
ADANSONIA, sér. 3 • 2011 • 33 (2)
Nomenclatural changes and taxonomic notes in the tribe Morindeae (Rubiaceae)
APPENDIX 1 – Continuation.
Species recognized in this study
Morinda pedunculata Valeton
Morinda persicifolia Buch.-Ham.
Morinda pumila Craib
Morinda ramosa (Lauterb.) Razafim.
& B.Bremer
Morinda rosiflora Y.Z.Ruan
Morinda royoc L.
Homotypic synonyms
Heterotypic synonyms
Morinda lanceolata Wall.; M. talmyi
Pierre ex Pit.
Sarcocephalus ramosus Lauterb.;
Sarcopygme ramosa (Lauterb.)
Setch. & Christoph.
Morinda ferruginea A.Rich.;
M. yucatanensis Greenm.
Morinda scabrida Craib
Morinda sessiliflora Bertol.
Morinda titanophylla E.M.A.Petit
Morinda tomentosa B.Heine ex Roth
Morinda turbacensis Kunth
Morinda undulata Y.Z.Ruan
Siphonandrium intricatum K.Schum.
ADANSONIA, sér. 3 • 2011 • 33 (2)
309