Reinstatement of Gymnosporia (Celastraceae): implications for the Flora Malesiana region
ABSTRACT 2003. Reinstatement of Gymnosporia (Celastraceae): implications for the Flora Malesiana region. Telopea 10(1): 155–167. The reinstatement of the genus Gymnosporia, comprising all the spiny members previously included in the genus Maytenus, has implications for the Flora Malesiana region. An account is given of six species and two varieties in the region (Gymnosporia curtisii, G. diversifolia, G. inermis, G. littoralis, G. nitida, G. spinosa var. spinosa and var. parva). The new combination G. littoralis (Backer) Jordaan is made (based on Gymnosporia montana var. littoralis) and a neotype and lectotype designated for two other taxa. G. emarginata from India and Sri Lanka is also included since this name was previously misapplied to plants in the Flora Malesiana region. Maytenus rapakir is mentioned as probably belonging to Gymnosporia. Maytenus cupularis, which has a racemose inflorescence, is related to the Australian species of Maytenus s. lat. and is retained in Maytenus.
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ABSTRACT: The phylogeny of Celastraceae tribe Celastreae, which includes about 350 species of trees and shrubs in 15 genera, was inferred in a simultaneous analysis of morphological characters together with nuclear (ITS and 26S rDNA) and plastid (matK, trnL-F) genes. A strong correlation was found between the geography of the species sampled and their inferred relationships. Species of Maytenus and Gymnosporia from different regions were resolved as polyphyletic groups. Maytenus was resolved in three lineages (New World, African, and Austral-Pacific), while Gymnosporia was resolved in two lineages (New World and Old World). Putterlickia was resolved as nested within the Old World Gymnosporia. Catha edulis (qat, khat) was resolved as sister to the clade of Allocassine, Cassine, Lauridia, and Maurocenia. Gymnosporia cassinoides, which is reportedly chewed as a stimulant in the Canary Islands, was resolved as a derived member of Gymnosporia and is more closely related to Lydenburgia and Putterlickia than it is to Catha. Therefore, all eight of these genera are candidates for containing cathinone- and/or cathine-related alkaloids.Molecular Phylogenetics and Evolution 06/2008; 48(2):745-57. · 4.02 Impact Factor
Reinstatement of Gymnosporia (Celastraceae):
implications for the Flora Malesiana region
Marie Jordaan and A.E. van Wyk
Jordaan, Marie1and van Wyk, A.E.2 (1National Botanical Institute, Private Bag X101, Pretoria, Republic
of South Africa 0001; e-mail: firstname.lastname@example.org; 2Department of Botany, University of Pretoria,
Pretoria, Republic of South Africa 0002; e-mail: email@example.com) 2003. Reinstatement of
Gymnosporia (Celastraceae): implications for the Flora Malesiana region. Telopea 10(1): 155–167. The
reinstatement of the genus Gymnosporia, comprising all the spiny members previously included in
the genus Maytenus, has implications for the Flora Malesiana region. An account is given of six
species and two varieties in the region (Gymnosporia curtisii, G. diversifolia, G. inermis, G. littoralis,
G. nitida, G. spinosa var. spinosa and var. parva). The new combination G. littoralis (Backer) Jordaan
is made (based on Gymnosporia montana var. littoralis) and a neotype and lectotype designated for
two other taxa. G. emarginata from India and Sri Lanka is also included since this name was
previously misapplied to plants in the Flora Malesiana region. Maytenus rapakir is mentioned as
probably belonging to Gymnosporia. Maytenus cupularis, which has a racemose inflorescence, is
related to the Australian species of Maytenus s. lat. and is retained in Maytenus.
The Celastraceae account for the Flora Malesiana was published by Ding Hou in 1962
and 1964. His genus concepts coincide with those of Exell (1952, 1953), Blakelock
(1956), Marais (1960) and Robson (1965, 1966 & 1994) in that he considers Gymnosporia
and Maytenus to be congeneric. New research on the family suggests that larger
genera need to be recircumscribed into smaller segregate ones. The genus Gymnosporia
(Wight & Arn.) Hook.f. has recently been reinstated (Jordaan & Van Wyk 1999, Archer
& Jordaan 2000) to include all its spiny members, previously placed under Maytenus
Molina sens. lat. A study of Gymnosporia (the southern African species) was
undertaken and the subdivision of the genus into eight sections was proposed
(Jordaan 1995), based mainly on fruit and seed morphology as well as leaf anatomy; a
monographic study covering the genus on a worldwide basis is currently in progress.
All the Gymnosporia species in the Flora Malesiana region have seed with a basal aril
and are placed in G. section Tenuispina M.Jordaan ined. Diagnostic characters of
Gymnoporia include the presence of brachyblasts and spines, leaves which are in
fascicles or alternate, the inflorescence which is a dichasium, the flowers which are
mostly functionally unisexual, the fruit which is a dehiscent capsule and the seeds
which have an aril.
Gymnosporia is an Old World genus of suffrutices, shrubs and trees. It comprises about
108 members, occurring in the whole of Africa, Madagascar and adjacent islands,
southern Spain, the near Middle East, Afghanistan, Pakistan, India, Sri Lanka,
Thailand, Vietnam, S China, Taiwan, Ryukyu Islands (Japan), Malesia, and Queensland
(Australia), with G. vitiensis (A.Gray) Seem. endemic to the Polynesian Islands.
The genus has two main centres of diversity on the Afro-Arabian continent: (i) NE
tropical Africa and tropical Arabia and (ii) southern Africa. There is a high degree of
regional endemism at both specific and sectional level and there are only a few
widespread species in the genus. Five members of Gymnosporia occur on the
Philippines, which is the highest level of species diversity in the Flora Malesiana
region. The purpose of this contribution is to provide an account of the members of
Gymnosporia native to the Flora Malesiana region.
Key to the species
1a Plants with spines of two kinds: axillary ones and lateral branches ending in a spine:
2a Capsules 2-valved:
3a Leaves usually smaller than 30 mm; China, Taiwan, Ruykyu Islands and N Philippines
.................................................................................................................... 6. G. diversifolia
3b Leaves usually longer than 30 mm; Thailand, Malaysia and Java .......... 5. G. littoralis
2b Capsules 3-valved:
4a Plants totally glabrous; India and Sri Lanka .......................................... 7. G. emarginata
4b Plants with puberulous or muricate branches; Thailand, Malaysia and Java ..................
.................................................................................................................................. 5. G. littoralis
1b Plants usually without spines or when spines present then only axillary ones, lateral branches
not ending in a spine:
5a Pericarp of capsules thick (± 1.5 mm), woody .................................................... 4. G. nitida
5b Pericarp of capsules thin, papery (less than 1 mm):
6a Leaf apex acute to short-acuminate; Thailand and Malaysia ...................... 3. G. curtisii
6b Leaf apex obtuse, rounded to emarginate; Taiwan, Philippines, Indonesia, New Guinea,
7a Brachyblasts usually present; leaves coriaceous; capsules longer than 6 mm
...................................................................................................................... 1. G. inermis
7b Brachyblasts usually absent; leaves membranous; capsules shorter than 6 mm
...................................................................................................................... 2. G. spinosa
1. Gymnosporia inermis Merrill & Perry
(Merrill & Perry 1939: 335).
Type: Papua New Guinea: Western Division: Tarara, Wassi Kussa River, Brass 8690, Jan
1937; holo A; iso L!, LAE!, US!
Gymnosporia montana sensu Benth.
(Bentham 1863: 400), non M.A.Lawson (M.A.Lawson 1875: 621).
Maytenus emarginata sensu Ding Hou
(Ding Hou 1962: 241), quoad specimens from N Queensland, Philippines, Celebes,
Moluccas and New Guinea; sensu Jessup (Jessup 1984: 164).
Scandent shrub or small tree, up to 5 m high, glabrous in all parts, with spines when
young, becoming spineless with age; brachyblasts well developed on older branches;
branches angular when young, grey, with inconspicuous lenticels. Leaves alternate on
young branchlets, fasciculate on older ones, ovate, elliptic or oblong, (30–)40–110(–140)
Telopea 10(1): 2003
× (15–)20–76(–110) mm, apex rotundate to retuse, or subacute to emarginate, base
cuneate to rotund, margin distinctly crenulate-serrate, reticulate venation prominent
on both sides; petioles up to 12 mm long, sometimes pinkish-tinged. Inflorescence
axillary, solitary or few-flowered in short cymes; peduncle ± 10 mm long, pink-tinged;
pedicels up to 4 mm long, pink-tinged, bracts lanceolate. Flowers white or cream-
coloured, 5–7(–9) mm in diameter, fragrant. Sepals semi-orbiculate. Petals oblong-
ovate, 2–3(–4) × 1–2 mm, apex obtuse, margin fimbriate. Male flowers: stamens slightly
shorter than petals; pistillode short; stigmas absent. Disc green, 5-lobed. Female
flowers: staminodes shorter than stamens of male flowers: ovary 3-locular, green; style
as long as ovary, 3-branched. Capsule 3-valved, obconic-trigonous, 3-angled, 6–12 mm
long, green becoming red. Seed ellipsoid, glossy, 3.5–5.5 × 2–4 mm; aril reddish,
reduced to a basal rim. (Fig. 1).
Diagnostic characters: plants when young with spines, but spines not developing on
older stems; well developed brachyblasts present; leaves coriaceous and losing spines
with age; well developed brachyblasts present; leaves coriaceous and usually longer
than 40 mm, apex rounded, obtuse to emarginate or subacute; capsules 3-valved, red;
seed with a basal aril.
Distribution and ecology: (Celebes and Moluccas) Indonesia, New Guinea, East Coast
of the Philippines, Taiwan and Queensland (Australia). Growing at elevations from
near sea level and locally abundant on the coast on sand at the edge of mangrove
forest, or secondary forest, at forest margins, in thickets on beaches and hillsides and
on sea cliffs, often on limestone.
Selected specimens examined: INDONESIA: Kei Island (Kai Kecil), Jensen 80, 25 Mar 1922 (GH, L);
Tanimbar Islands: Selaru, Namtabung, Van Borssum 3176, 17 Mar 1956 (K, L); Moluccas: Sula
Islands, Tandjong Baliha, Bloembergen 4389, 28 July 1939 (GH, SING); Aru Islands: Pulau Trangan,
Kp. Kerei, Turner 115, 17 Apr 1993 (GH, L). Sulawesi (Celebes): Tukangbesi, Binongko, Elbert 2559,
23 July 1909 (L).
NEW GUINEA: West Papua: Radjah Ampat, Waigeo Island, Lupintol on SW coast of Majalibit Bay,
Van Royen 5454, 8 Feb 1955, (L); Merauke, Koch Expedition 1904–1905 (L).
Jordaan and Wyk, Reinstatement of Gymnosporia
Fig. 1. Gymnosporia inermis. Sulawesi (Celebes): Binongko, Pulau, Elbert 2559 (L).
Papua New Guinea: Tarara, Wassi Kussa River, Brass 8690, Jan 1937 (L, LAE, US); Lea Lea, NW of
Pt. Moresby, Pullen 3359, 19 July 1962 (CANB, LAE); Daru Island, Brass 6229 29 Mar 1936 (L); Nanuk
Island, East New Britian Prov., Nagari UPNG 7303, 3 Nov 1981 (LAE).
PHILIPPINES: Dalupiri Island: Babuyan Group, 19°10'N 121°15'E, Bartlett 15075, 15168, 31 Oct–
5 Nov 1935 (GH); Luzon: Isabela Prov: Palanan municipality, Dipoduanaw Point, 17°06.8'N
122° 31.1'E, Leonardo Co 3393, 3394, 18 Apr 1991 (GH, K, L, PNH); Luzon: Sierra Madre Mountains
of Dingalan, sea level, 15° 29' N 121° 23' E, Jacobs 7772, 16 Mar 1968 (L).
AUSTRALIA: Queensland: Jack Jacky Ck, Cape York, 10° 58' S 142° 30' E, Jones 3821, 25 Oct 1965
(CANB); Portland Roads, 12°35' S 143°25' E, Dockrill 468, 4 July 1972 (BRI, K); Cairns: Webb & Tracey
8193, June 1965 (BRI, CANB).
2. Gymnosporia spinosa (Blanco) Merrill & Rolfe
(Merrill & Rolfe 1908: 109); (Merrill 1918: 235); (Loesener 1942: 151).
Cupania spinosa Blanco
(Blanco 1837: 184).
Type: Philippines: Luzon Island: Bulacan Prov., Angat, Merrill: Species Blancoanae 349,
Sep 1913; neo US! (designated here); isoneo GH!, L!
Selected specimens examined: PHILIPPINES: Luzon: Cagayan Prov.: Nabanagan, Pacheco 29523,
Mar 1923 (GH); Bulacan Prov.: Angat, Merrill: Species Blancoanae 349, Sep 1913 (L, US); Rizal Prov.:
San Juan del Monte, Pr Manila, Vidal 189 (GH); Lubang Islands: Merrill 978, Apr 1903 (GH); Panay:
Port Iloilo, Vidal 189, Nov–Dec 1958 (GH).
var. parva Merrill & Rolfe
(Merrill & Rolfe 1908: 110).
Type: Philippines: Luzon Island: Rizal Province, Montalban, Mariquina River, Merrill
5070 March 1905; holo PNH†; iso K!, L!, NY! US!
G. philippinensis Vidal (Vidal 1885: 103), nom. nudum quoad specimen Cuming 1575
(L). Rolfe (1886) stated that this is just a manuscript name and not validly published.
Selected specimens examined: PHILIPPINES: Luzon: Cuming 1575 (L); Rizal Prov., Montalban,
Elmer 12573, Jan 1910 (GH, L); Merrill 5070, Mar 1905 (K, L, NY, US).
Plants sometimes with small thin axillary spines, even on fruiting branches. Adistinct
species with two varieties, which can be separated from G. inermis by axillary spines
more often present, its membranous leaves (20–35 × 8–20 mm) which are narrower,
more elliptic and thinner. It has 3-valved capsules like G. inermis, but the capsules are
smaller, shorter than 5 mm. The difference between the two varieties lies in the size of
the leaves; var. parva has smaller leaves, shorter and narrower than 20 mm. (Fig. 2).
Distribution: apparently endemic to the Philippines.
Notes: Merrill and Rolfe (1908) stated, and we agree, that G. spinosa is not the same as
G. montana (Roth ex Roem. & Schult.) M.A.Lawson from India, which is the type
species of the genus Gymnosporia.
Merrill and Perry (1939) stated that G. inermis is closely related to G. spinosa but lacks
the spines of the latter species. G. inermis and G. spinosa are indeed very closely related.
Specimens of G. spinosa var. spinosa and var. parva have often short axillary spines, lack
brachyblasts, their leaves are thinner in texture and the reticulate venation on both
surfaces of the leaves is more prominently raised than in G. inermis; their capsules are
smaller (usually less than 6 mm long).
Telopea 10(1): 2003
Blanco (1837) originally described G. spinosa in Flora de Filipinas, but apparently there
is no original material in PNH of Blanco’s names of new taxa. Merrill’s specimen at US
is therefore selected here as a neotype, following the recommendation of Nicolson and
3. Gymnosporia curtisii King
(King 1896: 353); (Prain 1904: 198); (Ridley 1922: 451).
Type: Malaysia: Kedah Prov., Polo Tanyury Piri, Curtis 2500, Sep 1890; holo K!
Maytenus curtisii (King) Ding Hou
(Ding Hou 1962: 240).
Illustration: Ding Hou (1962: fig. 2a–d).
Erect or scandent shrub or small tree; spines robust, in axils of leaves. Leaves
coriaceous, elliptic, sometimes broadly elliptic, 62–85 (–110) × 27–36 (–90) mm, base
cuneate, tapering into petiole, apex acute to short-acuminate, rarely obtuse, margin
crenulate, principal lateral veins 7–9(–15), obliquely spreading towards margin and
curved upwards, reticulate venation obvious above; petiole 5–9 mm long.
Inflorescences cymose, usually crowded towards apex of brachyblasts, axillary,
puberulous when young; peduncle 5–10 mm long; bracts lanceolate, short-fimbriate;
pedicels 2–5 mm long. Flowers white. Sepals deltoid or semi-orbicular, 0.5–0.75 mm in
diameter, margins sparsely ciliate. Petals ovate or obovate-oblong, 2.5–3.2 × 1.5 mm,
obtuse, ± entire. Disc fleshy, rounded, 1.5–2.0 mm in diameter. Male flowers with
stamens ± 2.5 mm long; anthers broad-ovoid, 0.5 mm long, slightly apiculate;
pistillode with short style, stigmas absent. Female flowers with staminodes very short
or abortive; ovary trilocular; style longer than ovary; stigma 3-lobed. Capsule 3-valved,
obconic-trigonous, concave at tip, ± 15 × 13 mm. Seed ellipsoid, slightly irregularly
rugose, 7–8 × 4–5 mm; aril ± flat or shallowly disc-like and attached laterally at base.
Diagnostic characters: plants usually with axillary spines; leaf apex acute to short-
acuminate; capsule 15 mm long, 3-valved; seed with a flat or shallowly disc-like aril,
laterally attached at base.
Distribution and ecology: Thailand and Malaysia. It occurs on limestone at sea level,
or in lowland forests.
Jordaan and Wyk, Reinstatement of Gymnosporia
Fig. 2. Gymnosporia spinosa var. parva. Philippines: Rizal Province, Montalban, Elmer 12573 (L).