Nomenclatural Notes on Garcinia (Clusiaceae) from Madagascar and
the Comoros
Patrick W. Sweeney
University of Missouri–Saint Louis, One University Boulevard, St. Louis, Missouri
63121-4400, U.S.A.; Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri
63116-0299, U.S.A. Current address: Yale University Herbarium, Peabody Museum of
Natural History, 170 Whitney Avenue, New Haven, Connecticut 06511, U.S.A.
patrick.sweeney@yale.edu
Zachary S. Rogers
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63116-0299, U.S.A.
zachary.rogers@mobot.org
ABSTRACT . New results from phylogenetic analyses
utilizing chloroplast and nuclear DNA markers agree
with morphology in support of the unification of all of
Rheedia L. and part of Ochrocarpos Thouars with
Garcinia L. and show that species occurring in
Madagascar and the Comoros fall into four separate
lineages, which are designated here as informal
species groups. An examination of Garcinia from
these areas results in the recognition of 32 currently
described species, all but one of which are endemic.
The widespread African species, G. livingstonei T.
Anderson, is noted for the first time to occur in
Mayotte. Eleven new combinations are published
here: G. ambrensis (H. Perrier) P. Sweeney & Z. S.
Rogers, G. anjouanensis (H. Perrier) P. Sweeney & Z.
S. Rogers, G. arenicola (Jumelle & H. Perrier) P.
Sweeney & Z. S. Rogers, G. calcicola (Jumelle & H.
Perrier) P. Sweeney & Z. S. Rogers, G. dalleizettei (H.
Perrier) P. Sweeney & Z. S. Rogers, G. mangorensis
(R. Viguier & Humbert) P. Sweeney & Z. S. Rogers, G.
multifida (H. Perrier) P. Sweeney & Z. S. Rogers, G.
parvula (H. Perrier) P. Sweeney & Z. S. Rogers, G.
thouvenotii (H. Perrier) P. Sweeney & Z. S. Rogers, G.
tsaratananensis (H. Perrier) P. Sweeney & Z. S.
Rogers), and G. urschii (H. Perrier) P. Sweeney & Z. S.
Rogers. Three new names, G. dauphinensis P.
Sweeney & Z. S. Rogers, G. megistophylla P. Sweeney
& Z. S. Rogers, and G. tsimatimia P. Sweeney & Z. S.
Rogers, are provided for O. parvifolius Scott-Elliot, R.
megaphylla H. Perrier, and R. pedicellata (Jumelle &
H. Perrier) H. Perrier, respectively. Lectotypes are
designated for 12 names: G. crassiflora Jumelle & H.
Perrier, G. disepala Vesque, G. melleri Baker, G.
polyphlebia Baker, G. verrucosa Jumelle & H. Perrier,
O. ambrensis H. Perrier, O. macrophyllus O. Hoffmann, O. parvifolius, O. parvulus H. Perrier, O.
NOVON 18: 524–537. PUBLISHED
ON
tsaratananae H. Perrier, R. arenicola Jumelle & H.
Perrier, and R. calcicola Jumelle & H. Perrier.
Key words: Clusiaceae, Comoros, Garcinia, Guttiferae, Madagascar, Mammea, Ochrocarpos, Rheedia,
Tsimatimia, Xanthochymus.
When broadly circumscribed, the genus Garcinia L.
contains more than 250 species (Jones, 1980; Stevens,
2006) of mostly small- to medium-sized dioecious trees
and has a pantropical distribution with centers of
diversity in Madagascar and Southeast Asia. Garcinia
mangostana L., known by the common name mangosteen, is probably the most widely recognized member
of the genus, being a popular fruit tree from Southeast
Asia and the subject of a burgeoning herbal supplement
industry. The genus is notable for its high sympatric
species diversity (e.g., Whitmore, 1998; Lee et al.,
2002; Thomas et al., 2003) and the large amount of
morphological variation present in the flowers.
Almost since its inception, the limits of Garcinia
and related genera, including four with representatives in Madagascar, i.e., Ochrocarpos Thouars,
Rheedia L., Tsimatimia Jumelle & H. Perrier, and
Xanthochymus Roxburgh, have been debated (e.g.,
Planchon & Triana, 1860; Vesque, 1893; Engler,
1893, 1925; Jumelle & Perrier de la Bâthie, 1910;
Perrier de la Bâthie, 1948, 1951; Robson, 1958;
Adams, 1970; Jones, 1980; Gustafsson et al., 2002;
Stevens, 2006). In the last infrageneric classification
of the group, Jones (1980) adopted a broad concept of
the genus (including Ochrocarpos p.p., Rheedia,
Tsimatimia, and Xanthochymus) and recognized 14
sections that were delimited largely by staminate
floral morphology.
Recently, a broad-scale molecular phylogenetic
study of Garcinia utilizing two nuclear DNA markers
16 DECEMBER 2008.
doi: 10.3417/2006146
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Garcinia (Clusiaceae) from Madagascar
525
(granule-bound starch synthase [GBSSI] and ITS) has
been completed (Sweeney, 2008). The results of that
study, which include a geographically, taxonomically,
and morphologically comprehensive sampling of species, indicate that a broad circumscription of the genus
Garcinia is supported (sensu Jones, 1980; Stevens,
2006) and suggests that the genus has representatives
from four lineages in Madagascar and the Comoros.
Two of these lineages have taxa that are widely treated
as Garcinia, whereas the remaining two include
members that were placed into the genera Ochrocarpos
and Rheedia by Perrier de la Bâthie (1948, 1951).
Perrier de la Bâthie (1951) placed all Malagasy
Clusiaceae with unisexual flowers and fused sepals in
bud into Ochrocarpos. Kostermans (1956, 1961) and
de Wilde (1956) both commented on the affinity of
some species of Ochrocarpos to Mammea L., with de
Wilde (1956) suggesting that Ochrocarpos in its
entirety should be sunk into Mammea. While Kostermans (1956, 1961) thought that all of the Asian and
some of the Malagasy Ochrocarpos should be moved
into Mammea, he doubted that Malagasy Ochrocarpos
completely belonged there, suggesting instead that
Ochrocarpos, in part, should be maintained to include
those species of the genus with phalangiate androecia
and leaves lacking the higher order venation he
considered to be characteristic of true Mammea (see
below). Earlier, Vesque (1893: 482) regarded species of
Ochrocarpos as being closely related to Garcinia and
transferred O. decipiens Baillon into Garcinia. Jones
(1980) and Stevens (2005) agreed that Ochrocarpos
comprised two different groups of species, one related
to Garcinia and the other to Mammea. The Ochrocarpos
species related to Garcinia can be recognized by their
seeds, which possess an embryo with a grossly swollen
(vs. unswollen) hypocotyl and minute (vs. large)
cotyledons, by the stamens arranged in phalangiate
(vs. non-phalangiate or fasciculate) androecia in the
staminate flowers, and by the leaves without punctate
glands and with exudate-containing canals transversely
intersecting the secondary veins (vs. leaves with
punctate glands occupying the areoles and rarely with
exudate-containing canals). Minute cotyledons are less
than 1/10 of the length of the embryo, and large
cotyledons account for most of the embryo (Brandza,
1908; Stevens, 2006). The affinity of the Ochrocarpos
species possessing non-fasciculate androecia to Mammea has recently been supported by a combined
phylogenetic analysis of molecular and morphological
data (Notis, 2004), and the molecular phylogenetic
study of Sweeney (2008) supports the placement of the
Ochrocarpos species with phalangiate androecia in
Garcinia. Kostermans (1956, 1961) and Stevens (2005)
have transferred Malagasy species to Mammea, but four
Ochrocarpos species with phalangiate androecia treated
by Perrier de la Bâthie (1951) still lack valid names in
Garcinia. Perrier de la Bâthie (1948, 1951) recognized
G. cauliflora Baker, but Stevens (2005) transferred the
species to Mammea, creating the combination, M.
cauliflora (Baker) P. F. Stevens.
Earlier authors distinguished the genus Rheedia
from Garcinia by its flowers possessing two sepals
instead of four (e.g., Planchon & Triana, 1860; Engler,
1893, 1925). Robson (1958) correctly pointed out that
this distinction breaks down when one takes into
account the total variation within the two genera.
Robson (1958) and later Adams (1970) argued for the
inclusion of Rheedia in Garcinia, and this circumscription has been adopted in recent treatments (e.g.,
Kearns et al., 1998; Schatz, 2001). Molecular
phylogenetic studies support this view (Gustafsson,
2002; Sweeney, 2008). Robson (1958) noted that
Rheedia and Garcinia were published simultaneously
and that, when united, Garcinia should be the
preferred generic name in consideration of its size.
Ten species recognized as Rheedia by Perrier de la
Bâthie (1951) lack valid names in Garcinia.
Molecular phylogenetic data, strongly supported by
morphology, suggest that the Garcinia of Madagascar
and the Comoros belong to four different clades that
generally correspond to previously recognized taxonomic sections within Garcinia. Each of these clades
is made up of species that share unique combinations
of morphological characters (Sweeney, 2008). The 32
species recognized in the synopsis (Table 1) are
placed, using morphology, into one of four informal
groups that correspond to those clades identified in
the molecular analyses. While our groups roughly
correspond to previously recognized sections, we do
not assign species to formally named sections because
our groups do not absolutely correspond to any
previous author’s circumscription. These groups can
be identified with the accompanying key and
illustrations of the staminate and pistillate flowers of
representative species of all four groups (Figs. 1, 2).
Fourteen species lack names in the genus, thus
leading us to propose 11 new combinations and three
new names for 10 species in Rheedia and for four
species from Ochrocarpos.
The typification status of all names is addressed
following the protocol employed by Turland and Jarvis
(1997: 458–461). Our investigation is based on an
examination of herbarium specimens deposited at B,
BM, G, K, MO, P, TAN, and TEF.
A KEY TO THE FOUR SPECIES GROUPS OF GARCINIA IN MADAGASCAR
COMOROS
AND THE
1a. Staminate flowers with a disk occupying the center
of the flower; pistillate flowers with an annular or
lobed disk beneath the ovary.
526
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Table 1. The 32 species of Garcinia in Madagascar and the Comoros, their previous assignment in Perrier de la Bâthie
(1951), and the species group to which each is assigned here. The four species groups are recognized by morphology and
supported by evidence of the first author (Sweeney, 2008) utilizing the two nuclear DNA markers GBSSI and ITS.
Species group
Brindonia group
Paragarcinia group
Rheedia group
Accepted name
Previous assignment
Garcinia asterandra Jumelle & H. Perrier
G. chapelieri (Planchon & Triana) H. Perrier
G. crassiflora Jumelle & H. Perrier
Garcinia cerasifer (H. Perrier) P. F. Stevens
G. dauphinensis P. Sweeney & Z. S. Rogers
G. decipiens (Baillon) Vesque
G. disepala Vesque
G. madagascariensis (Planchon & Triana) Pierre
G. melleri Baker
G. multifida (H. Perrier) P. Sweeney & Z. S. Rogers
G. orthoclada Baker
G. parvula (H. Perrier) P. Sweeney & Z. S. Rogers
G. pauciflora Baker
G. polyphlebia Baker
G. tsaratananensis (H. Perrier) P. Sweeney & Z.
S. Rogers
Garcinia ambrensis (H. Perrier) P. Sweeney & Z.
S. Rogers
G. anjouanensis (H. Perrier) P. Sweeney & Z. S. Rogers
G. aphanophlebia Baker
G. arenicola (Jumelle & H. Perrier) P. Sweeney
& Z. S. Rogers
G. calcicola (Jumelle & H. Perrier) P. Sweeney
& Z. S. Rogers
G. commersonii (Planchon & Triana) Vesque
G. dalleizettei (H. Perrier) P. Sweeney & Z. S. Rogers
G. livingstonei T. Anderson
G. mangorensis (R. Viguier & Humbert) P. Sweeney
& Z. S. Rogers
G. megistophylla P. Sweeney & Z. S. Rogers
G. pervillei (Planchon & Triana) Vesque
G. thouvenotii (H. Perrier) P. Sweeney & Z. S. Rogers
G. tsimatimia P. Sweeney & Z. S. Rogers
G. urschii (H. Perrier) P. Sweeney & Z. S. Rogers
Xanthochymus group Garcinia capuronii Z. S. Rogers & P. Sweeney
G. lowryi Z. S. Rogers & P. Sweeney
G. verrucosa Jumelle & H. Perrier
2a. Stamens and staminodes free in staminate and
pistillate flowers . . . . Rheedia group (Fig. 1A–D)
2b. Stamens and staminodes phalangiate in staminate and pistillate flowers . . . . . . . . . . .
. . . . . . . . . . . Xanthochymus group (Fig. 1E, F)
1b. Staminate flowers lacking a disk and instead with a
mushroom-shaped pistillode or with stamens occupying the center of the flower; pistillate flowers
lacking a disk.
3a. Sepals 4 and free in bud; staminate flowers
lacking a well-developed pistillode and stamens free; ovary in pistillate flowers and fruit
with deep furrows down the septal radius . . . . . . . . . . . Brindonia group (Fig. 2A–E)
3b. Sepals usually 2 and fused in bud; staminate
flowers with a mushroom-shaped pistillode
and with stamens in phalanges; ovary in
pistillate flowers and fruit usually lacking
furrows down the septal radius . . . . . . . . . .
. . . . . . . . . . . . . Paragarcinia group (Fig. 2F–H)
Garcinia asterandra Jumelle & H. Perrier
G. chapelieri (Planchon & Triana) H. Perrier
G. crassiflora Jumelle & H. Perrier
Ochrocarpos cerasifer H. Perrier
O. parvifolius Scott-Elliot
O. decipiens Baillon
O. multiflorus O. Hoffmann
Rheedia madagascariensis Planchon & Triana
O. madagascariensis Choisy (non Planchon &
Triana)
O. multifidus H. Perrier
O. orthocladus (Baker) H. Perrier
O. parvulus H. Perrier
G. pauciflora Baker
G. chapelieri (Planchon & Triana) H. Perrier
O. tsaratananae H. Perrier
Rheedia ambrensis H. Perrier
R. anjouanensis H. Perrier
R. aphanophlebia (Baker) H. Perrier
R. arenicola Jumelle & H. Perrier
R. calcicola Jumelle & H. Perrier
R. madagascariensis (Planchon & Triana) H.
Perrier
R. dalleizettei H. Perrier
R. mangorensis R. Viguier & Humbert
R. megaphylla H. Perrier
R. pervillei Planchon & Triana
R. thouvenotii H. Perrier
R. pedicellata (Jumelle & H. Perrier) H. Perrier
R. urschii H. Perrier
G. verrucosa Jumelle & H. Perrier
DISCUSSION OF SPECIES GROUPS
RHEEDIA GROUP
Thirteen members of this group (Table 1; Fig. 1A–
D) were treated as Rheedia by Perrier de la Bâthie
(1951) and were placed into Jones’ (1980) invalidly
published Garcinia sect. Rheedia. The new report of G.
livingstonei T. Anderson from Mayotte material marks
the first instance the species has been found occurring
naturally outside of continental Africa. The 14 species
of this group in Madagascar and the Comoros can be
recognized by their staminate flowers having numerous
free stamens surrounding a disk located in the center of
the flower (or inserted on disk in G. livingstonei) and by
their pistillate flowers with an annular disk beneath the
ovary. The results of Sweeney (2008) suggest that the
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Garcinia (Clusiaceae) from Madagascar
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Figure 1. A–D. Rheedia group: Garcinia commersonii. —A, B. Staminate flowers, showing numerous free stamens and the
central disk. —C, D. Pistillate flowers, showing numerous free staminodes (disk not visible). E, F. Xanthochymus group: Garcinia
lowryi. —E. Pistillate flower, showing four antepetalous phalanges alternating with disk lobes (one sepal and two petals removed,
phalange in foreground broken). —F. Staminate flower, showing four antepetalous phalanges and the central disk (one petal in
foreground removed). Sources: A, B, G. Schatz et al. 3371 (MO); C, D, Service Forestier 22875 (MO); E, N. Dumetz & G. McPherson
1156 (MO); F, G. McPherson & N. Dumetz 14648 (TEF). A–D illustrated by J. Myers; E, F illustrated by L. R. Andriamiarisoa.
Malagasy and Comorian Rheedia and G. livingstonei are
sister taxa, and that clade is sister to a clade of South
American Rheedia. The entire Rheedia group plus the
G. livingstonei clade is nested within a group of African
Garcinia representing Garcinia sect. Rheediopsis Pierre
(sensu Jones, 1980).
Included species. Garcinia ambrensis (H. Perrier)
P. Sweeney & Z. S. Rogers, G. anjouanensis (H.
Perrier) P. Sweeney & Z. S. Rogers, G. aphanophlebia
Baker, G. arenicola (Jumelle & H. Perrier) P.
Sweeney & Z. S. Rogers, G. calcicola (Jumelle & H.
Perrier) P. Sweeney & Z. S. Rogers, G. commersonii
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Figure 2. A–E. Brindonia group: Garcinia chapelieri. —A–C. Staminate flowers, showing four sepals, four petals, and
numerous free stamens. —D, E. Pistillate flowers, showing several deep furrows down the septal radius of the ovary (E with two
sepals and one petal removed). F–H. Paragarcinia group: Garcinia decipiens. —F. Pistillate flower, showing four petals (sepals
not visible), several free stamens, the ovary, and the stigma. —G. Staminate flower bud, longitudinal section, showing fused
two sepals, two petals, two phalanges, and the mushroom-shaped pistillode. —H. Staminate flower, showing petals, four
antepetalous phalanges, and the pistillode. Sources: A–C, G. Schatz et al. 3373 (MO); D, E, P. Lowry et al. 4000 (MO); F, J.
Aridy & A. Moı̈se 183 (MO); G, H, Service Forestier 21601 (P). All parts illustrated by J. Myers.
(Planchon & Triana) Vesque, G. dalleizettei (H.
Perrier) P. Sweeney & Z. S. Rogers, G. livingstonei T.
Anderson, G. mangorensis (R. Viguier & Humbert) P.
Sweeney & Z. S. Rogers, G. megistophylla P.
Sweeney & Z. S. Rogers, G. pervillei (Planchon &
Triana) Vesque, G. thouvenotii (H. Perrier) P.
Sweeney & Z. S. Rogers, G. tsimatimia P. Sweeney
& Z. S. Rogers, G. urschii (H. Perrier) P. Sweeney &
Z. S. Rogers.
XANTHOCHYMUS GROUP
Until recently, this group (Table 1; Fig. 1E, F) was
represented in Perrier de la Bâthie (1951) by Garcinia
verrucosa Jumelle & H. Perrier, an endemic Malagasy
species, but two additional species were described by
Rogers and Sweeney (2007). The three species
representing this group in Madagascar are distinct
from other Malagasy and Comorian Garcinia by having
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Garcinia (Clusiaceae) from Madagascar
staminate flowers lacking a pistillode and possessing
four antepetalous phalanges that are composed of
incompletely fused filaments surrounding a disk in the
center of the flower and by their pistillate flowers with
a lobed disk beneath the ovary (lobes alternate with
the phalanges). Garcinia verrucosa is placed among a
monophyletic group of species from Garcinia sect.
Xanthochymus (sensu Jones, 1980) by molecular data
(Sweeney, 2008). This clade has species distributed
into Africa, Madagascar, India, Nepal, southern
China, and Malesia (Sweeney, 2008).
entirely of taxa endemic to Madagascar; however, the
exact position of this lineage within Garcinia remains
unresolved (Sweeney, 2008).
Included species. Garcinia capuronii Z. S. Rogers
& P. Sweeney, G. lowryi Z. S. Rogers & P. Sweeney, G.
verrucosa.
BRINDONIA GROUP
In Madagascar this group (Table 1; Fig. 2A–E)
contains three of the six species treated as Garcinia by
Perrier de la Bâthie (1951) and is distinct from the
other groups by having flowers with four sepals free in
bud and staminate flowers that possess numerous
stamens occupying the center of the flower (pistillode
and disk absent), and by ovaries and fruits usually
with deep furrows down the septal radius. Molecular
phylogenetic analyses (Sweeney, 2008) suggest that
these species fall within a group comprised almost
entirely of taxa previously placed into Garcinia sect.
Brindonia by Jones (1980).
Included species. Garcinia asterandra Jumelle &
H. Perrier, G. chapelieri (Planchon & Triana) H.
Perrier, G. crassiflora Jumelle & H. Perrier.
PARAGARCINIA GROUP
This strictly Malagasy group contains 12 species
(Table 1; Fig. 2F–H) and largely corresponds to
Garcinia sect. Paragarcinia (Baillon) Vesque sensu
Jones (1980). Eight species were previously recognized (Perrier de la Bâthie, 1951) in Ochrocarpos sect.
Paragarcinia Baillon, while a ninth was unplaced
within Ochrocarpos. Two species, G. pauciflora Baker
and G. polyphlebia Baker, were last treated under
Garcinia, and G. madagascariensis (Planchon &
Triana) Pierre was treated as a synonym of the invalid
combination Rheedia madagascariensis (Perrier de la
Bâthie, 1951). The Paragarcinia group is characterized by flowers with two sepals fused in bud (except G.
madagascariensis and G. pauciflora) and staminate
flowers with a mushroom-shaped pistillode and four to
eight or more antepetalous (occasionally branched)
phalanges of sessile to subsessile stamens. Phylogenetic analyses including species from this group
suggest that it is a strongly supported clade comprised
529
Included species. Garcinia cerasifer (H. Perrier) P.
F. Stevens, G. dauphinensis P. Sweeney & Z. S.
Rogers, G. decipiens (Baillon) Vesque, G. disepala
Vesque, G. madagascariensis, G. melleri Baker, G.
multifida (H. Perrier) P. Sweeney & Z. S. Rogers, G.
orthoclada Baker, G. parvula (H. Perrier) P. Sweeney
& Z. S. Rogers, G. pauciflora, G. polyphlebia, G.
tsaratananensis (H. Perrier) P. Sweeney & Z. S.
Rogers.
A SYNOPSIS OF GARCINIA IN MADAGASCAR AND THE COMOROS
Thirty-two species of Garcinia are treated in this
taxonomic synopsis (refer to Table 1 for species group
designation). Thirty are endemic to Madagascar, G.
anjouanensis is endemic to the Comoros, and G.
livingstonei occurs in Mayotte and Africa. One
dubious name, Ochrocarpos madagascariensis Choisy,
is also discussed. This alphabetically arranged list
includes 11 new combinations (G. ambrensis, G.
anjouanensis, G. arenicola, G. calcicola, G. dalleizettei,
G. mangorensis, G. multifida, G. parvula, G. thouvenotii, G. tsaratananensis, G. urschii) and three new
names (G. dauphinensis, G. megistophylla, G. tsimatimia). Twelve names are lectotypified. Garcinia
ochrocarpoides Jumelle & H. Perrier and O. madagascariensis were lectotypified by previous authors.
1. Garcinia ambrensis (H. Perrier) P. Sweeney & Z.
S. Rogers, comb. nov. Basionym: Rheedia ambrensis H. Perrier, Mém. Mus. Natl. Hist. Nat.,
n.s., 24: 91. 1948. TYPE: Madagascar. ‘‘Près du
lac Maudit, sur la Montagne d’Ambre, au N de la
Grande Ile…,’’ = fl., H. Perrier de la Bâthie 17730
(holotype, P030787; isotype, P030788).
2. Garcinia anjouanensis (H. Perrier) P. Sweeney
& Z. S. Rogers, comb. nov. Basionym: Rheedia
anjouanensis H. Perrier, Mém. Mus. Natl. Hist.
Nat., n.s., 24: 91. 1948. TYPE: Comoros.
‘‘Anjouan…,’’ ca. 700 m, fr., Lavanchie 20
(holotype, P030789; isotype, P030790).
3. Garcinia aphanophlebia Baker, J. Linn. Soc.,
Bot. 25: 295. 1889. Rheedia aphanophlebia
(Baker) H. Perrier, Mém. Mus. Natl. Hist. Nat.,
n.s., 24: 90. 1948. TYPE: Madagascar. ‘‘E. de la
grande Ile…,’’ ca. 500–1500 m, = fl., R. Baron
next 5797 (holotype, K000240255; isotypes,
K000240256, P030791).
The Kew sheet (K000240255) is the only relevant
material bearing the collection number (i.e., ‘‘next
5797’’) as it was cited in the protologue (Baker, 1889:
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Novon
295). On that sheet, someone anonymously marked
out the ‘‘next 5797’’ and wrote in ‘‘5796.’’ The other
sheet at K (K000240256) is only numbered with
‘‘5796,’’ whereas the final digit of the number on the P
sheet (P030791) was changed anonymously from ‘‘7’’ to
‘‘6.’’
la Bâthie 8148 were found at P (P030793, P030794).
These syntypes only bear fruits.
4. Garcinia arenicola (Jumelle & H. Perrier) P.
Sweeney & Z. S. Rogers, comb. nov. Basionym:
Rheedia arenicola Jumelle & H. Perrier, Ann.
Sci. Nat. Bot., sér. 9, 11: 269. 1910. TYPE:
Madagascar. ‘‘…région de Madirovalo, dans le
Boina, et près de Manongarivo, dans l’Ambongo…sablonneux secs du Bongo-Lava,’’ Oct.
1900, = fl., H. Perrier de la Bâthie 1119 bis
(lectotype, designated here, P030792).
Several unvouchered localities in northwestern
Madagascar were prominently mentioned in the
protologue of Garcinia arenicola (Jumelle & Perrier
de la Bâthie, 1910: 270). Three separate collections
(Perrier de la Bâthie 1119, 1119 bis, 1119 ter), each
represented by a single sheet in the P herbarium, bear
some part of the locality information from the
protologue. Two of these (1119 bis, 1119 ter) are
annotated as types of the basionym in Perrier de la
Bâthie’s own hand, and Perrier de la Bâthie 1119 bis
(P030792) is designated as the lectotype, as it is in the
best condition. The two other collections are regarded
as syntypes.
5. Garcinia asterandra Jumelle & H. Perrier, Ann.
Sci. Nat. Bot., sér. 9, 11: 280. 1910. TYPE:
Madagascar. ‘‘…Massif du Manongarivo…,’’
1400 m, = fl., H. Perrier de la Bâthie 5311
(holotype, P030763).
6. Garcinia calcicola (Jumelle & H. Perrier) P.
Sweeney & Z. S. Rogers, comb. nov. Basionym:
Rheedia calcicola Jumelle & H. Perrier, Ann.
Sci. Nat. Bot., sér. 9, 11: 266. 1910. TYPE:
Madagascar. ‘‘…dans la Moyenne-Mahavavy,…le Tompoketsa,…environs de Majunga,…
ravin d’Antsahobé…,’’ Oct. 1904, = fl., imm. fr.,
H. Perrier de la Bâthie 1752 (lectotype, designated here, P030795).
Several unvouchered localities were mentioned in
the protologue of Rheedia calcicola (Jumelle & Perrier
de la Bâthie, 1910: 268). These place names have
been traced to two collections (Perrier de la Bâthie
1752 and 8148) deposited in the P herbarium that
were annotated as types of the basionym in Perrier de
la Bâthie’s own hand. Sheet P030795 of Perrier de la
Bâthie 1752 bears staminate flowers and immature
fruit and, as the most complete collection, is
designated as the lectotype. Two sheets of Perrier de
7. Garcinia capuronii Z. S. Rogers & P. Sweeney,
Syst. Bot. 32: 773. 2007. TYPE: ‘‘Fianarantsoa,
Kianjavato, entre Ifanadiana et Anosivolo,’’ 6
Dec. 1964, R fl., fr., Service Forestier (Capuron)
23916 (holotype, P030776; isotypes, P030774,
P030775, P030777, TEF).
8. Garcinia cerasifer (H. Perrier) P. F. Stevens,
Harvard Pap. Bot. 9: 433. 2005. Basionym:
Ochrocarpos cerasifer H. Perrier, Mém. Mus.
Natl. Hist. Nat., n.s., 24: 103. 1948. Mammea
cerasifer (H. Perrier) Kostermans, Djawatan
Kehutanan Indonesia Bagian Planologi Kehutanan, Djalen Perniagaan 44: 12. 1956. TYPE:
Madagascar. ‘‘…Massif de l’Ikongo…,’’ ca.
1400 m, fr., R. Decary 5659 (holotype,
P030860; isotypes, K00240251, P00389082,
TAN000266).
9. Garcinia chapelieri (Planchon & Triana) H.
Perrier, Mém. Mus. Natl. Hist. Nat., n.s., 24: 97.
1948. Basionym: Ochrocarpos chapelieri Planchon & Triana, Ann. Sci. Nat. Bot., sér. 4, 14:
366. 1860. TYPE: Madagascar. s. loc., imm. fr.,
L. Chapelier s.n. (holotype, P030769).
Perrier de la Bâthie (1948, 1951) treated Garcinia
polyphlebia as a synonym of G. chapelieri. An
examination of the original material of G. polyphlebia
and G. chapelieri at K and P indicates that the two are
quite different morphologically and that G. polyphlebia is instead closer to the former Ochrocarpos species
of Garcinia (i.e., the Paragarcinia group herein). Given
these differences, we resurrect G. polyphlebia (treated
below). Newly collected pistillate material at MO
closely matches the type material of G. chapelieri
vegetatively, and the ovaries and fruits of this material
have several distinctive furrows down the septal
radius. Perrier de la Bâthie (1951: 60) described the
fruits of G. chapelieri as smooth; however, the fruit
description was probably based on the type material of
G. polyphlebia under the assumption that the two
species were synonymous.
10. Garcinia commersonii (Planchon & Triana)
Vesque, Monogr. Phan. 8: 484. 1893. Basionym:
Rheedia commersonii Planchon & Triana, Ann.
Sci. Nat. Bot., sér. 4, 14: 312. 1860. TYPE:
Madagascar. s. loc., R fl., P. Commerson s.n.
(holotype, P-JU 11865).
Garcinia pachyphylla Baker, J. Linn. Soc., Bot. 25: 295.
1889. TYPE: Madagascar. ‘‘North-west…,’’ = fl., R.
Baron 5757 (holotype, K000240238; isotypes,
K000240239, P030798).
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2008
Sweeney & Rogers
Garcinia (Clusiaceae) from Madagascar
Ochrocarpos humblotii Drake, Bull. Mens. Soc. Linn. Paris 2:
1220. 1896, as ‘‘Humbloti.’’ Rheedia humbloti (Drake)
R. Viguier & Humbert, Rev. Gén. Bot. 25, bis: 637.
1914. TYPE: Madagascar. s. loc., = fl., L. Humblot
391 (holotype, P030800; isotypes, K000240243,
K000240244, P030801, P030802).
13. Garcinia dauphinensis P. Sweeney & Z. S.
Rogers, nom. nov. Replaced name: Ochrocarpos
parvifolius Scott-Elliot, J. Linn. Soc., Bot. 29: 5.
1891. TYPE: Madagascar. ‘‘Woods near Fort
Dauphin…,’’ = fl., G. Scott-Elliot 2710 (lectotype, designated here, K000240237; isotypes,
P00568804, P030823).
Garcinia commersonii and its synonyms were
previously recognized as synonyms of the invalid
name ‘‘Rheedia madagascariensis (Planchon & Triana)
H. Perrier’’ by Perrier de la Bâthie (1948: 92, 1951:
46). See further discussion of the name under G.
madagascariensis (Planchon & Triana) Pierre. The
provenance in the protologue of the original material
(Humblot 391) for Ochrocarpos humblotii was cited as
‘‘Iles Comores, ou Madagascar?’’. However, Perrier de
la Bâthie (1948: 92) indicated that the type was
collected in the littoral forest of Madagascar, not the
Comoros, and the taxon has apparently not been
collected outside of Madagascar thus far.
11. Garcinia crassiflora Jumelle & H. Perrier,
Ann. Sci. Nat. Bot., sér. 9, 11: 279. 1910. TYPE:
Madagascar. ‘‘…dans le Manongarivo et le
Sambirano,’’ = fl., fr., H. Perrier de la Bâthie
5305 (lectotype, designated here, P030764;
isotype, P030765).
In the protologue, Jumelle and Perrier de la Bâthie
(1910: 280) cited the provenance of the original
material as ‘‘Manongarivo et le Sambirano.’’ Three
relevant collections, Perrier de la Bâthie 5305
(P030764, P030765), 5313 (P030767), and 5313 bis
(P030766), have been found at P, and all are annotated
as types with the place of publication of the protologue
in Perrier de la Bâthie’s own hand, and bear
inscriptions with at least one of the two place names
mentioned in the protologue. Perrier de la Bâthie 5305
(P030764) is the most complete specimen of the
original material and is designated as the lectotype.
The isolectotype consists of nothing more than a few
fruits glued to the sheet. Perrier de la Bâthie 5313 and
5313 bis should be regarded as syntypes.
12. Garcinia dalleizettei (H. Perrier) P. Sweeney &
Z. S. Rogers, comb. nov. Basionym: Rheedia
dalleizettei H. Perrier, Mém. Mus. Natl. Hist. Nat.,
n.s., 24: 93. 1948. TYPE: Madagascar. ‘‘EST: Mt.
Vatohazo, dans le bassin infèrieur du Mangoro…,’’ 300–400 m, = fl., H. Perrier de la Bâthie
18064 (holotype, P030796; isotype, P030797).
Two sheets of Perrier de la Bâthie 18064 have been
found at P. Sheet P030796 bears a lectotype sticker,
but this is an error as the collection is the only original
material cited in the protologue (Perrier de la Bâthie,
1948: 93).
531
Two collections, Scott-Elliot 2710 and 2840, both
noted to be collected near Fort Dauphin, were cited in
the protologue (Scott Elliot, 1891: 5). The former,
Scott-Elliot 2710, is the more complete collection, and
the K sheet (K000240237), clearly annotated as the
type specimen of Ochrocarpos parvifolius, is selected
as the lectotype. The epithet parvifolia is validly
occupied by Garcinia parvifolia (Miquel) Miquel
(Miquel, 1864) for a species native to Peninsular
Malaysia, Borneo, and Sumatra (Whitmore, 1973). We
choose our new epithet to refer to the collection
locality of the type specimen, but the species may
occur as far north as the Masoala Peninsula.
14. Garcinia decipiens (Baillon) Vesque, Monogr.
Phan. 8: 482. 1893. Basionym: Ochrocarpos
decipiens Baillon, Adansonia 11: 370. 1876.
TYPE: Madagascar. ‘‘Nossi-Bé, in humidis…,’’
Jan. 1841, = fl., A. Pervillé 421 (holotype,
P030815; isotypes, K000240241, P030816).
Ochrocarpos macrophyllus O. Hoffmann, Sert. Pl. Madagasc.
7. 1881. TYPE: Madagascar. ‘‘Nossi-komba,’’ Dec.
1879, = fl., J. Hildebrandt 3239 (lectotype, designated here, P030818; isotypes, BM, K000240245,
G00090054, P030817, P00462369).
Garcinia ochrocarpoides Jumelle & H. Perrier, Ann. Sci. Nat.
Bot., sér. 9, 11: 275. 1910. Ochrocarpos jumellei R.
Viguier & Humbert, Rev. Gén. Bot. 25, bis: 635. 1914,
nom. illeg. superfl. TYPE: Madagascar. ‘‘Dans le massif
du Manongarivo…bois du versant du Sambirano…,’’ R
fl., H. Perrier de la Bâthie 5312 (lectotype, designated
by Perrier de la Bâthie, 1948: 109, P00462373).
The original material of Ochrocarpos macrophyllus
(Hildebrandt 3239) deposited at B is no longer extant
(R. Vogt, pers. comm.). Two sheets of original material
are present at P, and the most complete (P030818) is
designated as the lectotype.
Ochrocarpos jumellei R. Viguier & Humbert is
illegitimate and superfluous according to Article 52.1
of the International Code of Botanical Nomenclature
(McNeill et al., 2006: 92–93) because Viguier and
Humbert (1914: 635) did not adopt the available
epithet of the clearly indicated basionym when
publishing the name.
15. Garcinia disepala Vesque, Epharmosis, part 2:
pl. 159. 1889. Ochrocarpos multiflorus O.
Hoffmann, Sert. Pl. Madagasc. 7. 1881. TYPE:
532
Novon
Madagascar. ‘‘Nossi-komba, arbor ad littora
maris floribus lacteis…,’’ Feb. 1880, = fl.,
J. Hildebrandt 3337 (lectotype, designated
here, G00090050 [stamp 724413]; isotypes,
G00090049 [2], G00090050 [stamp 724412],
K00380200, P030820, P030821).
Ochrocarpos multiflorus and Garcinia disepala are
homotypic, the former having been published about
eight years before the latter, but the epithet already
existed in Garcinia for the validly published G.
multiflora Champion ex Bentham (1851), a name that
pertains to a widespread Asian species occurring in
southern China, Taiwan, and northern Vietnam (Li et
al., 2007).
The original material of Hildebrandt 3337 deposited at B is no longer extant (R. Vogt, pers. comm.), so
the sheet at G bearing the accession number
G00090050 and stamped with 724413 is designated
as the lectotype.
16. Garcinia livingstonei T. Anderson, J. Linn.
Soc., Bot. 9: 263. 1867. TYPE: Africa. ‘‘Hab. in
rupibus schistosis prope flumen Zambesi, in
horto botanico Calcuttensi ex Africa culta,’’ R fl.,
D. Kirk s.n. (holotype, CAL not seen).
17. Garcinia lowryi Z. S. Rogers & P. Sweeney,
Syst. Bot. 32: 775. 2007. TYPE: Madagascar.
‘‘Toliara, Fort-Dauphin, W of town in forest
called Mandena, trail through Botanical Garden,’’ 6 Dec. 1989, = fl., G. McPherson & N.
Dumetz 14648 (holotype, MO; isotypes, P, TAN,
TEF).
18. Garcinia madagascariensis (Planchon & Triana) Pierre, Fl. Forest. Cochich. 1(5): 5. 1883.
Basionym: Xanthochymus madagascariensis
Planchon & Triana, Ann. Sci. Nat. Bot., sér. 4,
14: 305. 1860, as ‘‘Xanthochymus ? madagascariensis.’’ TYPE: Madagascar. s. loc., = fl., P.
Commerson s.n. (holotype, P-JU 11879; isotype,
P0303771).
The history of Garcinia madagascariensis and its
basionym, Xanthochymus madagascariensis, has been
confused in the literature and herbarium almost since
the names were first published. The name ‘‘Xanthochymus ? madagascariensis’’ first appeared in Planchon and Triana (1860: 305) and was reproduced two
years later in a reprint entitled Mémoire sur la famille
des Guttifères (Planchon & Triana, 1862: 150).
Baillon (1877: 402) suggested that Xanthochymus
should be united with Garcinia and erroneously cited
page 303 of Planchon and Triana’s publication. Pierre
(1883) formally transferred the species to Garcinia,
attributing authorship of the name incorrectly to
Baillon, presumably because Baillon (1877) listed
‘‘Xanthochymus madagascariensis Pl. et Triana. Mém
Guttif. p. 150. Habite Madagascar (Commerson)’’ as
the basionym of G. madagascariensis.
Perrier de la Bâthie (1948: 97) referred the name
Garcinia madagascariensis (with ‘‘Baillon ex Pierre’’
as the author) to an entity known to him only as an
incomplete, unnumbered Commerson specimen
housed at P (no accession number). In that publication, he went on to mention ‘‘Xanthochymus madagascariensis Pl. et Tri., Mem. Ternstr. Gutt., 150’’ in
synonymy, which was an incorrect reference to the
reprinted protologue appearing in the aforementioned
Mémoire sur la famille des Guttifères (Planchon &
Triana, 1862: 150). While Perrier de la Bâthie (1948:
97) did not provide a description of ‘‘G. madagascariensis Baillon ex Pierre,’’ the species is noted in his
key of Garcinia as having ‘‘Un gros rudiment
styliforme de gynécée, coiffé d’un large stigmate
rouge…,’’ which could be accommodated within the
description provided in the protologue of X. madagascariensis. In this instance, it seems that Perrier de
la Bâthie (1948) recognized a taxon based on the type
of X. madagascariensis, and thus referred to the same
taxon recognized as such by Pierre (1883) and
Planchon and Triana (1860). Curiously, only two
years later Perrier de la Bâthie (1951) no longer
recognized the name G. madagascariensis, referring to
the name only as a synonym of his invalid combination, ‘‘R. madagascariensis.’’
Perrier de la Bâthie (1948: 92), under the genus
Rheedia, created a new invalid combination, R.
madagascariensis, which he also puzzlingly based on
Xanthochymus madagascariensis Planchon & Triana,
this time correctly citing ‘‘Ann Sc. Nat. Bot., sér. 4,
XIV (1860), 305’’ for the protologue of the basionym
(Planchon & Triana, 1860: 305). He also cited
‘‘Garcinia madagascariensis H. Bn., Hist. Pl., VI,
402’’ as a synonym, providing the same erroneous
place of publication and authorship as before. Perrier
de la Bâthie (1948: 92) synonymized several names
(G. commersonii, G. pachyphylla, Ochrocarpos humblotii, R. commersonii, and R. humblotii) with R.
madagascariensis. All of these taxa are morphologically similar in having staminate flowers with free
stamens arranged in a ring around a central disk, a
characteristic of our Rheedia group. The original
material of R. commersonii, like that of X. madagascariensis, is an unnumbered Commerson specimen,
and this similarity apparently led Perrier de la Bâthie
to mistakenly conclude that R. commersonii and X.
madagascariensis were homotypic. However, this is
extremely unlikely as the protologue of X. madagascariensis describes a plant quite different from that of
the original material of R. commersonii. Our study
suggests that the entity that Perrier de la Bâthie
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2008
Sweeney & Rogers
Garcinia (Clusiaceae) from Madagascar
(1948, 1951) referred to as ‘‘Rheedia madagascariensis’’ (nom. inval.) represents a distinct taxon and
consequently needs a valid name in Garcinia. Aside
from X. madagascariensis, we also agree that the
names Perrier de la Bâthie (1948, 1951) placed into
synonymy under R. madagascariensis refer to the
same taxon. The name R. commersonii Planchon
& Triana has priority when recognized as Rheedia,
and G. commersonii (Planchon & Triana) Vesque is
the correct name when Rheedia and Garcinia are
united.
In regard to the original material of Xanthochymus
madagascariensis, Planchon and Triana (1860: 305)
mentioned one or more unnumbered Commerson
collections in the protologue with the provenance
statement, ‘‘Commerson in herb. A. L. de Jussieu et in
herb. Thoüin, nunc Cambessèdes.’’ In the Jussieu
herbarium at P, we found a specimen attributed to
Commerson (sheet 11879) that corresponds to the
description in the protologue. This material consists of
a branch with leaves and staminate flowers, and we
consider it to represent holotype material overlooked
most recently by Perrier de la Bâthie (1948, 1951).
Another sheet in the general herbarium closely
matches the P-JU specimen and presumably corresponds to the collection cited in the protologue that
was originally housed separately as part of Thoüin’s
herbarium. We regard this specimen (P0303771) as
an isotype duplicate. Both sheets closely resemble
specimens subsequently collected from the southeast
coast of Madagascar, an area where Commerson was
known to have collected (Dorr, 1997: 93–95). This
material may have also served as the basis for plate
348 labeled as Garcinia madagascariensis (without
associated authorship or text) in Grandidier’s Atlas
(Drake, 1896).
When attempting to provide a name in Garcinia for
the combination Rheedia macrophylla (Martius) Planchon & Triana (Planchon & Triana, 1860), Verdcourt
(1976: 262) published the illegitimate, superfluous
new name G. megaphylla Verdcourt. He was apparently unaware that the combination was based on G.
macrophylla Martius (Martius, 1841), basing his new
name on ‘‘Rheedia macrophylla Planchon & Triana,’’
rather than on the actual basionym (for a more
complete discussion, see Hammel, 1989). Because the
epithet ‘‘megaphylla’’ is validly occupied in Garcinia,
a new epithet is needed for R. megaphylla when it is
transferred to Garcinia. In keeping with the meaning
conveyed in the replaced name and in reference to the
very large leaves found in this species (perhaps the
largest in the genus), we select the epithet ‘‘megistophylla,’’ meaning very big leaves.
19. Garcinia mangorensis (R. Viguier & Humbert)
P. Sweeney & Z. S. Rogers, comb. nov.
Basionym: Rheedia mangorensis R. Viguier &
Humbert, Bull. Soc. Bot. France 61: 131. 1914.
TYPE: Madagascar. ‘‘…bords de la Sahamarirana entre Ampasimpotsy et Bevalanirano, province d’Andovoranto, district de Moramanga…,’’
ca. 900 m, 24 Oct. 1912, R fl., R. Viguier & H.
Humbert 1011 (holotype, P030803; isotype,
G00018840).
20. Garcinia megistophylla P. Sweeney & Z. S.
Rogers, nom. nov. Replaced name: Rheedia
megaphylla H. Perrier, Mém. Mus. Natl. Hist.
Nat., n.s., 24: 90. 1948. TYPE: Madagascar. ‘‘Est
(S), forêt orientale,…sur le Mt. Vatovavy à ouest
de Mananjary…,’’ ca. 400–500 m, = fl., R.
Decary 13689 (holotype, P030804; isotype,
P030805).
533
21. Garcinia melleri Baker, J. Linn. Soc., Bot. 20:
92. 1883. TYPE: Madagascar. ‘‘Between Tamatave and Antananarivo…,’’ R fl., C. Meller s.n.
(lectotype, designated here, K000240242; isotype, P030819).
Ochrocarpos goudotianus Planchon & Triana, Ann. Sci. Nat.
Bot., sér. 4, 14: 365. 1860. TYPE: Madagascar. s. loc.,
1830, imm. fr., J. Goudot s.n. (holotype, P00568801;
isotype, G00090054).
Garcinia comorensis Drake, Bull. Mens. Soc. Linn. Paris 2:
1221. 1896. Ochrocarpos comorensis (Drake) R. Viguier
& Humbert, Rev. Gén. Bot. 25, bis: 634. 1914. TYPE:
Madagascar. Lac de Nossi-Vé, 11 Apr. 1882, R fl. bud,
L. Humblot 144 (holotype, P; isotype, K000240247).
Garcinia melleri was previously recognized as a
synonym of the name Ochrocarpos madagascariensis
Choisy (with the incorrect authorship ‘‘DC.’’) in
Perrier de la Bâthie (1948, 1951). However, the name
O. madagascariensis is dubious in its application and
is therefore treated as such at the end of the paper.
Two collections, Baron s.n. (K000380201) and
Meller s.n. (K000240242, P030819), were cited as
original material for Garcinia melleri. Earlier statements by Perrier de la Bâthie (1948: 110, 1951: 92)
mentioning Meller s.n. as the type of the name cannot
be considered an effective lectotypification, since a
depository was not specified and none of the material
at K or P bears any indication that Perrier de la Bâthie
examined it. Thus, it is impossible to distinguish
between the duplicates deposited in those herbaria,
and the K sheet of Meller s.n. is now designated as the
lectotype of the name because it is the most complete
specimen.
The provenance of the type material of Garcinia
comorensis was cited as ‘‘Iles Comores’’ in the
protologue, but the notes for Humblot 144 in the field
book register at P provide contradictory information,
534
namely ‘‘Garcinia comorensis Drake, est daté du 11
avril 1882, Lac de Nossi-Vé. Grand arbre 20 m.
Fleurs blanc crème, terre sablonneuse, 150 m. d’alt.’’
No other collections of G. melleri have been made
from the Comoros, so we infer that the protologue’s
provenance was probably an inadvertent mistake by
Drake.
22. Garcinia multifida (H. Perrier) P. Sweeney &
Z. S. Rogers, comb. nov. Basionym: Ochrocarpos
multifidus H. Perrier, Mém. Mus. Natl. Hist.
Nat., n.s., 24: 106. 1948. TYPE: Madagascar. ‘‘E.
du Lac Aloatra à Menaloha…,’’ = fl., G. Cours
672 (holotype, P030822).
23. Garcinia orthoclada Baker, J. Linn. Soc., Bot.
22: 446. 1887. Ochrocarpos orthocladus (Baker)
H. Perrier, Mém. Mus. Natl. Hist. Nat., n.s., 24:
108. 1948. TYPE: Madagascar. s. loc., fr., R.
Baron 3633 (holotype, K000240240; isotype,
P00462365).
Rheedia laka R. Viguier & Humbert, Bull. Soc. Bot. France
61: 131. 1914. TYPE: Madagascar. ‘‘…forêt d’Analamazoatra,’’ ca. 1000 m, 3 Nov. 1912, fr., R. Viguier &
H. Humbert 1111 (holotype, P00462367; isotypes,
B100158974, G00090051 p.p.).
Ochrocarpos ambrensis H. Perrier, Mém. Mus. Natl. Hist.
Nat., n.s., 24: 107. 1948, syn. nov. TYPE: Madagascar.
‘‘Forêt de la Montagne d’Ambre,…sur roches volcaniques, près de Diégo-Suarez (Nord)…,’’ ca. 1200 m, =
fl., H. Perrier de la Bâthie 17548 (lectotype, designated
here, P030814; isotypes, K000240257, P00568806,
TAN000265).
The isotype of Rheedia laka at G (G00090051) is
fragmentary, and the thick leafless branch on the left
hand side of the sheet is excluded as it does not
appear to represent the species.
Ochrocarpos ambrensis was described as a distinct
species by Perrier de la Bâthie (1948: 107) with some
reservations regarding its distinctness from O. orthocladus (; Garcinia orthoclada, sensu Baker). We
cannot find morphological differences to retain this
taxon as distinct from G. orthoclada and treat it here
for the first time as a new synonym of that name. Two
collections (Perrier de la Bâthie 17548, 18892), both
noted as coming from Montagne d’Ambre, were cited
in the protologue of O. ambrensis (Perrier de la Bâthie,
1948: 107). Perrier de la Bâthie 17548 (P030814) has
the most intact staminate flowers and is designated as
lectotype. The other syntype, Perrier de la Bâthie
18892 (P00568805), is in fruit.
24. Garcinia parvula (H. Perrier) P. Sweeney & Z.
S. Rogers, comb. nov. Basionym: Ochrocarpos
parvulus H. Perrier, Mém. Mus. Natl. Hist. Nat.,
n.s., 24: 108. 1948. TYPE: Madagascar. ‘‘…sur
le massif du Tsaratanana et ses abords, Nord du
Novon
Domaine central,…Sahandrakoto, au S. du
Tsaratanana…,’’ ca. 1500–2400 m, = fl., H.
Perrier de la Bâthie 16179 (lectotype, designated
here, P00462372; isotype, P00568803).
Two collections, Decary 1059 (P) and Perrier de la
Bâthie 16179 (P [2]), were cited in the protologue
(Perrier de la Bâthie, 1948: 108). The more complete
P sheet (P00568803) of the latter collection is
designated as the lectotype.
25. Garcinia pauciflora Baker, J. Linn. Soc., Bot.
20: 92. 1883. TYPE: Madagascar. ‘‘Forest of
Analamazaotra…,’’ imm. fr., R. Baron 1382
(holotype, K000240235; isotypes, K000240236,
P030779).
Garcinia cernua Baker, J. Linn. Soc., Bot. 22: 446. 1887.
TYPE: Madagascar. s. loc., imm. fr., R. Baron 2653
(holotype, K000240234; isotypes, K000240233,
P030780).
Perrier de la Bâthie (1948: 96) invalidly published
the infraspecific name ‘‘Garcinia pauciflora var.
depauperata’’ without an accompanying Latin description. The name, based on Viguier & Humbert 982,
refers to an entity he considered to be closely related
to G. pauciflora. However, it is clear after comparing
the type material with Perrier de la Bâthie’s (1948,
1951) descriptions of the two taxa in question that
they are not closely related because the invalid variety
depauperata has the staminate flower morphology of
the Brindonia group, whereas G. pauciflora belongs to
the Paragarcinia group.
26. Garcinia pervillei (Planchon & Triana) Vesque, Monogr. Phan. 8: 485. 1893, as ‘‘Garcinia ?
Pervillei.’’ Basionym: Rheedia pervillei Planchon
& Triana, Ann. Sci. Nat. Bot., sér. 4, 14: 312.
1860. Tsimatimia pervillei (Planchon & Triana)
Jumelle & H. Perrier, Ann. Sci. Nat. Bot., sér. 9,
11: 256. 1910. TYPE: Madagascar. ‘‘Île Nossibé…,’’ 18 Jan. 1841, imm. fr., A. Pervillé
407 (holotype, P030807; isotypes, P030808,
P030809).
The genus Tsimatimia was created by Jumelle and
Perrier de la Bâthie (1910: 263–264) to accommodate
Garcinia pervillei and T. pedicellata Jumelle & H.
Perrier (recognized below as G. tsimatimia), which
they believed differed from related genera in a few
characters of the calyx, androecium, and ovary.
Perrier de la Bâthie (1948) himself later placed
Tsimatimia in synonymy with Rheedia, based partly
on the observation that both species had free stamens
surrounding a disk. Tsimatimia has not been used by
subsequent authors.
Volume 18, Number 4
2008
Sweeney & Rogers
Garcinia (Clusiaceae) from Madagascar
535
27. Garcinia polyphlebia Baker, J. Linn. Soc.,
Bot. 22: 447. 1887. TYPE: Madagascar. Central
Madagascar, imm. fr., R. Baron 3101 (lectotype,
designated here, K000240249).
Natl. Hist. Nat., n.s., 24: 90. 1948. TYPE:
Madagascar. ‘‘…des terrains primitives gneissiques des forêts de l’Ankaizina…,’’ 1908, = fl., H.
Perrier de la Bâthie 5316 (holotype, P030806).
Two collections, Baron 3064 (K000240248,
K000240250,
P030768)
and
Baron
3101
(K000240249), were cited in the protologue of
Garcinia polyphlebia. The K sheet of Baron 3101 is
designated as the lectotype because it is the only
original element bearing the designation of type in
what appears to be Baker’s handwriting.
The pedicellata epithet is already in use in the genus
for Garcinia pedicellata (G. Forster) Seemann, an
endemic New Caledonian species (Smith, 1981; Jaffré
et al., 2004). Thus, we establish the new name, G.
tsimatimia, here for the Malagasy species. The epithet is
based on Tsimatimia, the generic name of the basionym,
which in turn was roughly based on the Malagasy
common name ‘‘Tsimatimanota,’’ meaning literally ‘‘not
punished for sinning’’ in the Malagasy language. Our
epithet is composed arbitrarily as supported by Art.
23.2 of the ICBN (McNeill et al., 2006: 45).
28. Garcinia thouvenotii (H. Perrier) P. Sweeney
& Z. S. Rogers, comb. nov. Basionym: Rheedia
thouvenotii H. Perrier, Mém. Mus. Natl. Hist.
Nat., n.s., 24: 90. 1948, as ‘‘Rheedia ? Thouvenotii.’’ TYPE: Madagascar. ‘‘Centre (E), forêt
d’Analamazoatra…,’’ 1000 m, = fl., E. Thouvenot s.n. (holotype, P030810; isotypes, P030811,
P030812).
29. Garcinia tsaratananensis (H. Perrier) P. Sweeney & Z. S. Rogers, comb. nov. Basionym:
Ochrocarpos tsaratananae H. Perrier, Mém. Mus.
Natl. Hist. Nat., n.s., 24: 108. 1948. TYPE:
Madagascar. ‘‘Sylve à lichens et forét à mousses,…dans le massif du Tsaratanana,’’ ca.
2000 m, = fl., H. Perrier de la Bâthie 16263
(lectotype, designated here, P00462368).
Two collections, Perrier de la Bâthie 16262
(P00568807) and 16263 (P00462368), both from the
Tsaratanana massif, were referred to indirectly by the
provenance given in the protologue of Ochrocarpos
tsaratananensis (Perrier de la Bâthie, 1948: 108). The
latter collection number is flowering and is designated
as the lectotype. The other syntype is in fruit. The
epithet as originally provided was derived from a
geographic locality and, according to Recommendation 60.D.1 of the ICBN (McNeill et al., 2006: 112),
should take the form of an adjective, and thus the
suffix of the epithet is adjusted here.
The name ‘‘Ochrocarpos tsaratananae var. rotundifolius’’ first appeared in Perrier de la Bâthie (1951:
86) followed by the phrase ‘‘Vig. et Humb., mss. in
Herb. Mus. Paris,’’ and a diagnosis and discussion in
French without the necessary validating Latin.
Characters cited as diagnostic in the protologue for
this variety were based on two specimens and break
down when additional collections are examined.
30. Garcinia tsimatimia P. Sweeney & Z. S.
Rogers, nom. nov. Replaced name: Tsimatimia
pedicellata Jumelle & H. Perrier, Ann. Sci. Nat.
Bot., sér. 9, 11: 265. 1910. Rheedia pedicellata
(Jumelle & H. Perrier) H. Perrier, Mém. Mus.
31. Garcinia urschii (H. Perrier) P. Sweeney & Z.
S. Rogers, comb. nov. Basionym: Rheedia urschii
H. Perrier, Mém. Mus. Natl. Hist. Nat., n.s., 24:
89. 1948. TYPE: Madagascar. ‘‘Centre (E), forêt
d’Analamazoatra…,’’ 1000 m, = fl., E. Ursch 42
(holotype, P030813).
32. Garcinia verrucosa Jumelle & H. Perrier, Ann.
Sci. Nat. Bot., sér. 9, 11: 277. 1910. TYPE:
Madagascar. ‘‘Sambirano,…bords du Ramena,’’
300 m, Aug. 1908, = fl., H. Perrier de la Bâthie
5314 (lectotype, designated here, P030782).
Four infraspecific taxa (subspecies ‘‘orientalis’’ and
‘‘typica,’’ and varieties ‘‘apiculata’’ and ‘‘piriformis’’)
were invalidly described under Garcinia verrucosa in
Perrier de la Bâthie (1948: 95–96), all of which lack a
validating Latin diagnosis or description. No original
material was explicitly cited in the protologue of G.
verrucosa (Jumelle & Perrier de la Bâthie, 1910: 277),
thus a collection with staminate flowers cited as
subspecies typica by Perrier de la Bâthie (1948) is
chosen for the lectotype of the species. An examination of herbarium material of G. verrucosa collected
subsequent to Perrier de la Bâthie’s work indicates
that the cited differences between his infraspecific
taxa break down.
DOUBTFUL NAME
Ochrocarpos madagascariensis Choisy, Prodr. 1: 560.
1824, non Ochrocarpos madagascariensis Planchon & Triana, Ann. Sci. Nat. Bot., sér. 4, 14:
364. 1860. TYPE: Madagascar. s. loc., pl. 26 in
Petit-Thouars, Hist. Vég. Isles Austral. Afriq.
1806 (lectotype, designated by Sprague, 1934:
89, pl. 26, Petit-Thouars, 1806).
Petit-Thouars first described the genus Ochrocarpos
in his Genera Nova Madagascariensia (1805: 15)
536
Novon
without any validating description of a species,
apparently basing the name on one of his own
personal herbarium collections. Choisy (1824: 560)
validated Petit-Thouars’ generic description by providing a species description of O. madagascariensis,
which he indicated as being based on an unnumbered
Petit-Thouars specimen. Original material, which
should have been deposited at P or possibly G-DC,
was not found in either herbarium. Planchon and
Triana (1860: 364) provided an amplified species
description of O. madagascariensis, based on a
different, unnumbered Petit-Thouars specimen at P,
which we refer to the invalid ‘‘Ochrocarpos planchonianus’’ (discussed below).
Sprague (1934: 89), having been unable to locate
any herbarium specimens traceable to Petit-Thouars,
provided a detailed discussion of the history of
Ochrocarpos and O. madagascariensis, and concluded that the only available original material of the
taxon was plate 26 in the very rare ‘‘edition 2’’ of
Histoire Végétaux Recueillis dans les Isles Australes
d’Afrique (Petit-Thouars, 1806). Sprague reproduced
the plate (labeled as Ochrocarpos and lacking
accompanying text) in his publication and clearly
indicated that the illustration should be regarded as
the type. The plant figured in the illustration is an
ample fruiting branch along with dissected young
fruits and does not conflict with either Petit-Thouars’
or Choisy’s descriptions. However, both descriptions
are so broad that specimens of many other species of
Malagasy Garcinia would also not obviously conflict
with either one, and the illustration is not identifiable to species.
Perrier de la Bâthie (1948: 103, 109) believed he
had rediscovered the missing Petit-Thouars original
material of the name at P, and annotated the sheet (no
accession number) with the inscription, ‘‘Type très
probable du genre Ochrocarpos Thouars et type de O.
madagascariensis DC.’’ The specimen is in poor
condition, consisting of five detached leaves, and is
unlikely to have been original material for Choisy’s
name. The specimen was not annotated as O.
madagascariensis by anyone prior to Perrier de la
Bâthie, and it obviously conflicts with the morphology
of the plant figured in plate 26 of Petit-Thouars
(1806). Leaves on the P sheet annotated by Perrier de
la Bâthie are obovate and rounded or emarginate at
their apex, while those figured in the plate of O.
madagascariensis are elliptic with an acute apex. The
leaves on this particular P sheet closely resemble
those of G. melleri in shape, venation pattern, and
color. Unfortunately, we have not found any material
approximating Sprague’s lectotype.
Perrier de la Bâthie (1948: 102) considered the
sterile Petit-Thouars material on which Planchon and
Triana (1860: 364) based their description of Ochrocarpos madagascariensis to be different from that which
Choisy had used, leading him to publish the invalid
name ‘‘Ochrocarpos ? Planchonianus’’ based on that
same material and without the necessary Latin
description or diagnosis. Two years later, Perrier de
la Bâthie (1951: 92), beneath O. madagascariensis
sensu Planchon & Triana (non O. madagascariensis
sensu Choisy), treated the presumed original material
as a young shoot or seedling of Rheedia, but he made no
mention of the name O. planchonianus. The PetitThouars material is still extant in the general P
herbarium (no accession number), and we agree that
it most likely represents immature vegetation of an
unidentifiable species belonging to the Rheedia group.
The authorship of Ochrocarpos madagascariensis has
been incorrectly ascribed to de Candolle in all relevant
literature since the name was first published in the
Prodromus, despite the obvious footnote at the bottom
of the first page of the Clusiaceae treatment indicating
that Choisy authored the work (Choisy, 1824: 557–
564). Botanists should be aware that several other
Clusiaceae names published in the Prodromus pertaining to a number of other geographical areas have been
erroneously attributed to de Candolle (for examples,
consult the International Plant Names Index website,
,http://www.ipni.org/index.html.).
Acknowledgments. The authors thank Robert Vogt
(B), Peter Stafford (BM), Laura Pearce and Yvette
Harvey (K), Peter Phillipson (MO), and Laurent Gautier
and Muriel Hecquet (G) for providing collection
assistance and high-resolution specimen images of
original material; Gerard Aymonin and Frédéric
Tronchet (P), and Nicholas Hind (K) for bibliographic
assistance; Anne-Elizabeth Wolf (P) for suggestions on
the manuscript; Jean-Nöel Labat (P) for making recent
collections of Mayotte material available; Petra Hoffmann (K) and Victoria C. Hollowell (MO) for helpful
review comments; John Myers and L. Roger Andriamiarisoa (MO) for providing illustrations; and Nicholas
Turland (MO) for insightful nomenclatural advice.
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