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au Bulletin du Muséum national d’Histoire naturelle,

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J. A. Doyle, University of California, Davis, USA

P.K. Endress, Institute of Systematic Botany, Zurich, Suisse

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adansonia

Éditions scientifiques du Muséum, Paris

Index des nouveautés taxonomiques et nomenclaturales

publiées dans Adansonia 20 (I), 1998

Landiopsis Capuron ex Bosser, gen. nov ... 132

Landiopsis capuronii Bosser, sp. nov ..... 132

Melaleuca brevisepala (j.W. Dawson) Craven & J.W. Dawson, corn b. nov. . 193

Melaleuca buseana (Guillaumin) Craven & J.W. Dawson, comb. nov. . 192

Melaleuca daivsonii Craven, nom. nov. ... 192

Melaleuca pancheri (Brongn. & Gris) Craven & J.W. Dawson, comb. nov. . 192

Melaleuca spbaerodendra Craven &C J.W. Dawson, nom. nov. . 192

Melaleuca sphaerodendra mai. microphylla (Virot) Craven & J.W. Dawson, comb. nov. . 193

Sylvichadsia Du Puy & Labat, gen. nov. . 165

Sylvichadsia grandidieri (Baill.) Du Puy & Labar, comb. nov . 168

Sylvichadsia grandifolia (R. Vig.) Du Puy & Labat, comb. nov. . 166

Sylvichadsia macrophylla (R. Vig.) Du Puy & Labat, comb. nov . 170

Sylvichadsiaperrieri (R. Vig.) Du Puy & Labat, comb. nov . 170

Weinmannia coodei H.C. Hopldns, sp. nov .52

Weinmannia croftii H.C. Hopkins, sp. nov. ,.76

Weinmannia devogelii H.C. Hopkins, sp. nov .. 48

Weinmannia eymaeana H.C. Hopldns, sp, nov .... 50

Weinmannia sect. Fasciculata Bernardi ex Hoogland & H.C. Hopkins, sect. nov. ..... 21

Weinmanniafurfuracea H.C. Hopkins, sp. nov . 49

Weinmannia hooglandü H.C. Hopkins & J.C. Bradford, sp. nov . 37

Weinmannia maxquesana var. myrsinites (Fosberg & Sachet) H.C. Hopkins & J. Florence, comb. nov.

.122

Weinmannia ouaiemensis (Guillaumin & Virot) Hoogland, comb. nov . 102

Weinmannia tremuloides H.C. Hopkins & J. Florence, sp. nov . 123

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Congress Street, Salem, Massachusetts 01970.

A révision of Weinmannia (Cunoniaceae)

in Malesia and the Pacific. 1. Introduction and

an account of the species of Western Malesia,

the Lesser Sunda Islands and the Moluccas

Helen C.F. HOPKINS

Laboratoire de Phanérogamie, Muséum national d’Histoire naturelle,

16 rue Buffon, 75005 Paris, France.

Masons Arms, Hutton Roof, via Carnforth, Lancashire, LA6 2PE, U.K.

With the collaboration of J.C. BRADFORD

Missouri Botanical Garden, P.O. Box 299,

St Louis, MO 63166. U S A.,

and Washington Universîty, St Louis, U.S.A.

KEYWORDS

Weinmannia,

Cunoniaceae,

inflorescence structure,

Malesia,

Pacific,

MOTS CLÉS

Weinmannia ,

Cunoniaceae,

structure inflorescenriellc,

Malésie,

Pacifique.

ABSTRACT

About 40 species of the l.irgcly tropical, montane genus Weinmannia occur

in Malesia and the Pacific belonging to two sections. Section Fasciculata is

largely Malcsian, extending eastwards to Fiji, and sect. Leiospemium is largely

Pacific, occurring as far west as Papua New Guînea. The structure of the

inflorescence provides previously unrecognised characters of taxonomie

importance, cspecially al the sectional lcvel. Because of the high levcls of

local endemism, species accounts foi the whole région are divided into four

parts. Descriptions, illustrations and distribution maps are provided here for

four species: W. Jraxinca, a widespread ochlospecies; W, ckmensiae, a local

endemic on ultramafic substrates in Sabah; W. apbanoneura , from Bornéo

and Sumatra, and W. hooglandii H.C. Hopkins & J.C. Bradford, sp. nov.,

from Peninsular Malaysia.

RÉSUMÉ

La Malésie et le Pacifique renferment environ 40 espèces de Weinmannia

(genre essentiellement tropical et montagneux), réparties dans deux sections :

sect. Fasciculata, présente surtout en Malésie et répandue il l’est jusqu à Fiji ;

sect. Leiospermum, surtout Pacifique, s'étendant à l’ouest jusqu'à la

Papouasie-Nouvelle-Guinée. La structure des inflorescences fournit des

caractères d importance taxonomique, surtout pour la distinction des sec¬

tions, qui n'avaient pas été utilisés jusqu’à maintenant. En raison de l'impor¬

tance de l’endémisme local, l'étude des espèces répandues dans l’ensemble de

cette région est divisée en quatre parties. Des descriptions, illustrations et

cartes de répartition sont fournies ici pour quatre espèces : W. Jraxinea une

ochloespèce largement répandue ; W. clemensiae endémique de Sabah, sur

substrats ultrarnafiques ; W. apbanoneura, de Bornéo et Sumatra, et W. hoo¬

glandii H.C. Hopkins & J.C. Bradford, sp. nov., de la Péninsule malaise.

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

INTRODUCTION

Weimnannia is the largest and most widespread

genus in the predominantly woody, Southern

hemisphere family, Cunoniaceae, and it contains

almost half the 300 or so species of the family as

it is currently circumscribed in BrummiïT

(1992). There are about 80 species in tropical

America (BERNARDI 1961, 1963a), ar least 30 in

Madagascar and rhe Mascarene Islands

(Bernardi 1964, 1965; J.C. Bradford pers,

comm.) and ca. 4() in Malesia and rhe Pacific

(BERNARDI 1964 and this révision). The présent

day distribution of the genus is thus rather

curious since it is absent from Africa, mainland

Asia excepr for Peninsular Malaysia, and from

Australia, which is part of the région of greatest

generic diversiry for the family. However, fossil

leaves with affinities to Weinmannia

(Weinmanniapkyllum) hâve been described from

Oligocène deposits in Tasmania (CARPENTF.R &

Buchanan 1993) suggesting thar it may once

hâve occurred in places where it is now extinct.

Weimnannia is largely tropical and montane

though it occurs in subtropical and temperate

forest in South America and in New Zealand, and

in lowland tropical forest in Madagascar. The

taxonomie history of the genus was discussed by

Bernardi (1961).

In the most recent treatment of the Malesian-

Pacific speçies, BERNARDI (1964) reçognised 40

species in two sections but he rarely gave com¬

plété descriptions except for new taxa. In add¬

ition, much new material has become available

since his work (e.g. 2/3 of the collections of W

urdanetensis Elmer and W. pullei Schltr. from the

Highlands of New Guinea hâve been collected

since 1959). R.D. HOOGIAND, who had a long-

standing monographie interest in the family,

started to révisé Weimnannia fot Malesia and the

Pacific in the 1970s. The présent study has

drawn on his extensive collection of taxonomie

literature, notes on morphology and typification,

and a catd index of specimens. However, the spe¬

cies concepts and descriptions are mine unless

otherwisc indicated.

My study was based largely on herbarium mat¬

erial with field observations in Malaysia, Fiji and

New Caledonia. It is divided into four parts: this

paper présents hackground information, an

account of the widespread and variable Wein-

manma fraxinen (D. Don) Miq. and of the

remaining species in Western Malesia, rhe fesser

Sunda Islands and the Moluccas; part 2

(HOPKINS 1998a) deals' with the species of

Sulawesi and the Philippines; part 3 (HOPKINS

199Sb) describes the species thar occur in New

Guinea and the islands of rhe western Pacific;

and part 4 (HOPKINS & FLORENCE 1998) deals

with the islands of the central Pacific and pré¬

sents an index to taxa, including synonyms.

Where appropriate, régional papers hâve been

further divided by island group, each with a

separate key. This arrangement bas been adopted

in part to reflect the high degree of régional and

island-group endemism, m part to reflect author-

ship of rhe accounts, and in pan because the prob-

lems remaining in Samoa and rhe Cook Islands

preclude a uniform treatmenr of ail taxa in a

single alphabetical list. l'hose wishing to see a

comprehensive key and alphabetical treatment of

species for Malesia will find this eventually in

Flora Malesiana, In the meanfime, the scctional

keys of BERNARD! (1964) are helpful. with sonie

minor adjustments for new and recircumstribed

taxa. An identification list of specimens seen

(excluding New Zealand, Samoa and the Cook

Islands) will bc deposited in major herbatia or

available from P.

Only one species, Weinmannia fraxinea, is

widespread in Malesia and the western Pacific,

occurring from northern Sumatra to the

Solomon Islands. It is described in some detail in

this paper. Where this and other species occur in

more thao one région (e.g. W. urdanetensis in the

Philippines and New Guinea; W. txtgua A.C.

Sm. in the Solomon Islands and Fiji), the des¬

cription and synonomy are given only once.

My species concept is morphological (see

McDADI 1995) and aims ro be consistent with

the définition of CttONQUIST (1978) as the small-

esr group that is consistently and persistently dis¬

tinct and distinguishable by oïdinary means. [t has

been suggested that over reliante on leaf chatacters

(especiaJIy size and number of leaflets) has lead to

taxonomie confusion and the over description of

species in Weinmannia. However, there are few

other characters available, as features of the flowers

ADANSONIA. sér. 3 • 1998 • 20(1)

Weinmannia in western Malesia

and fruits are rarely taxonomicaJIy uscful at the

species level. Thus many taxa differ by combina¬

tions of quantitative characters, none of which is

diagnostic on its own, The structure of the inflor¬

escence has provided some new characters, part-

icularly at the sectional level, but it is rather

uniform amongst the species in Malesia.

There has been a wealth of anatomical work on

Cunoniaceae and its relatives (e.g. DlCKISON

1975a,b, 1977. 1980a,b, 1984; Govn & Saxena

1976; Hideux & Ferguson 1976; Rao 8c

Dickison 1985; Rutishauser 8t Dickison

1989) and many of the characters investigated

hâve proved uscful in understanding relation-

ships wïthin the family and with its close relat¬

ives (Hufford 8c Dickison 1992). However,

species délimitation in Weinmannia is aggravated

by lack of strong morphological divergence, and

anatomical characters are unlikely to be useful at

this taxonomie level.

Within Cunoniaceae, Weinmannia is cjosely

related to Cunonia (ENGLER 1930-, HUFFORD &

Dickison 1992), with which it is sympatric in

New Caledonia. It differs from Cunonia in the

form of the floral dise (which is adnate to the

base of the ovary in Cunonia and composed of

either free lobes or a continuous ring, attached

only at its base, in Weinmannia), the dehiscence

of the fruit (dehiscing in a circle round the base

and acropetally along rhe sutures, the valves

remaining attached to the central column at the

apex in Cunorua\ dehiscing septicidally or septi-

fragally from the apex, and the valves and central

column remaining attached at the base in

Weinmannia) and in the seeds (angular and often

narrowly winged in Cunonia ; ellipsoidal, comose

at both ends or throughout and not winged in

Weinmannia) HUFFORD 8c DlCKlSON’s analysis

also suggests a close affinity with Pancheria but

this genux differs markedly from the other two in

its capitate inflorescences.

Weinmamiia has litrle économie importance in

Malesia and the Pacific. The bark contains high

levels of tannins and some species arc used for

tanning and firewood but few local uses are

recorded. The Flowers of Weinmannia racemosa

L. f. are visited by bees which in turn produce

honey on a commercial scale in New Zealand

(J.C. BRADFORD pers. comm.).

ECOLOGY AND DISTRIBUTION

Weinmannia species are typically trees and

shrubs of humid, montane forests, often growing

where there is persistent cloud cover and quite

high, well distributed rainfall. Twigs and leaf

blades are often covered with mosses and other

epiphytes. Within Malesia, the genus grows at

mid to high élévations (up to 2600-2800

(-3000) m) in Bornéo, Sulawcsi and New

Guinea) although a few species occur near sea

level. In the Pacific, Weinmannia is absent from

low coralline îslands but widespread and some-

times abundant on the high volcanic ones.

Végétation zones hère are more compressed than

in Malesia, and Weinmannia is typically found at

300-750+ m, The genus also occurs on islands of

continental origin, from 150-1550 m in New

Caledonia, and from ca. 300-1180 m in New

Zealand (W racem-osa , fide Wardle 1966).

Weinmannia often occurs on ridges, steep

slopes and in areas of disturbance due to tree

falls, landslides or volcanic éruptions. It is some-

times an early coloniser on volcanic slopes and

along roadsides (e.g. W. fraxinea in Sabah,

Weinmannia sp. in central Vin Levu) where it

tolérâtes high light intensity. While it is found

on a range of soil types, it often occurs on ones

with arypical minerai composition, including

soils derived from ulrramafic substrates and on

the leached soils associated with disturbed sites

in cloud fbrest. Frequent plant associâtes are the

fems Gleichenia and Dieranopteris which are typic-

al of burnt or repeatedly eut sites on impover-

ished soils, often rich in aluminium.

A few species hâve very spécifie écologies (e.g.

Weinmannia clemensiae Steenis from stunted

foresr on ultramafic soit) while others show

considérable ecological plasticity and are found

over a broad altitudinal range (e.g. W fraxinea in

parts of Malesia and W parviflora G. Forst in

Tahiti), and both these species arc morphologic-

ally vety variable. Weinmannia croftii H.C.

Hopkins from the Bismarck Archipelago is

found both as an early coloniser of volcanic

deposits and in adjacent closed, montane forest

(D. Frodin pers. comm.).

Most species of Malesian-Pacific Weinmannia

hâve quite restricted distributions and there is a

ADANSONIA, sér. 3 - 1998 • 20(1)

Weinmannia in western Malesia

high degree of island or island-group endemism.

Within Malesia, W. fraxinea is the only wide-

spread specics. In contrast with parts of the

Andes and Madagascar (J.C. BRADFORD pers.

comm.), there are fcw sympatric or co-occurring

species of Weinmannia in any part of its

Malesian-I’acific distribution It is of note that

there are only four species in New Caledonia,

which is a centre of species richness for the

Cunoniaceae, with a total of about 90 species. It

seems likely that here, Weinmannia is at least

partly replaced by Cunonia, which is morpholog-

ically sîmilar in many respects.

Weinmannia bas smal! seeds that are probably

wind-dispersed (see below). Anemochory, poss-

ibly with the aid of cyclones, and the opportuniste

ic ecological strategy of some species tie in well

with dispersai to, and establishment on, groups

of remote volcanic islands within the Pacific.

Despite rhe rôle that the break up of

Gondwanaland may hâve played in the world¬

wide distribution of the genus, more recent dis¬

persai events hâve dearly also had an impact.

Weinmannia is an important component of the

végétation in some parts of its distribution, .such

as the Society and Marquesas Islands, and New

Zealand. Those specics that, like W. Jraxinea, are

tolérant of natural disturbance, are likely to

become more abundant with increasing human

impact. However, some restricted endémies may

he of conservation interest including W. clemens-

iae from the Mt. Kinabalu massif in Sabah, W

paitensis Schltr., which is confjned to a few

mountains in cenrral and Southern New

Caledonia and W. ouaiemensis (Guillaumin &

Virot) Hoogland, Icnown from only a single local-

ity in north-eastem New Caledonia. Some other

species are known from very few collections but

we hâve too little information at présent to assess

their conservation status. Several species such as

W. rapensis F. Br. and W raiateensis J.W. Moore,

are endentic to small Pacific islands, but they are

not currently considered threatened.

RELATIONSHIPS OF THE

MALESIAN-PACIFIC SPECIES

Bernardi (1961, 1963a,b, 1964, 1965) pro-

ADANSONIA, sér, 3 ■ 1998 ■ 20(1)

posed an infrageneric classification of

Weinmannia with six sections. According to

BERNARDI, sections Weinmannia(e) and

Simplicifolia(e) are principally American, with a

few species in the Mascarene Islands (sect.

Weinmannia ); sections Spicala(e) and Inspersa(e)

are confined to the western Indian Océan

(Madagascar and Comores); and the Malesian-

Pacific species are placcd in sections

Fasctculata(e) and Leiospermum (referred to as

sect. Raccmosae by Bernardi). The lacter also

conrains two Malagasy species, W. comorcnsis 1 ul.

and W baehniana Bernardi, Sections Fasciculata

and Leiospermum are distinguished from the

others by the form of the floral dise, which is

composed of eight free lobes alternaung with rhe

filaments, rather than a continuous ring

(BERNARD! 1964: 132). In sect. Fasciculata , the

flowers are arranged in “pseudoracemes" whilst

in sect. Leiospermum they are dispersed. on pedi-

cels, in racemes or panicles (BERNARDI 1964:

132).

Prdiminary results of phylogenetîc studies in

progress, based on morphological characters,

especially of the inflorescence, and on DNA

sequeneing (BRADFORD in press and pers.

comm ) support the monophyly of both sections

Leiospermum (morphology; DNA still unresolv-

ed) and Fasciculata (strong support from DNA

sequences), although their relationships w r ith

other sections are still unclear. A few species were

misplaced by BERNARD!; the Indian Océan spe¬

cies he placée! in sect. Leiospermum belong with

the Malagasy sect. Spicata , and W descombesiana

Bernardi, placed by BERNARD! in sect.

Leiospermum, belongs with the Malesian sect.

Fasciculata (Bradford pers. comm.).

Section Fasciculata is a cohesive group of spe¬

cies. In addition to having the flowers inserted in

fasdcles on the inflorescence axis (to form the

“pseudoracemes" of Bernardi), these species

share a coinmon, rather complex, inflorescence

plan (see below). Weinmannia descombesiana is

somewhat problematical sitice it combines some

characters of both sections Fasciculata and

Leiospermum. Section Fasciculata is mo.stly

Malesian, extending as far as east as Fiji (Fig. I).

Section Leiospermum is confined to the Pacific,

extending westwards as far as the Bismarck

9 I

Hopkins H.C.F.

Archipelago and Karkar Island in Papua New

Guinea (Fig. 1). Most Pacific species form a clos-

ely knit group (e.g. W. parviflora, W. purpurea

L.M. Perry, W. vitiensis Seem.). Those from New

Caledonia form anocher group with minor différ¬

ences in the flowers, which are aJways bisexuaJ,

and the inflorescence structure. The New

Zealand species form a third group. Their flow¬

ers are inserted singly but hâve a greater tendancy

ro be perigynous and the structure of the inflor¬

escence shows some différences from the remain-

ing Pacific species. The species of rhe Marquesas

Islands, at th extreme east of Weiv-manmas dis¬

tribution in the Pacific, hâve their afflnities with

the other Pacific species, not the American ones,

NOTES ON MORPHOLOGY AND

BIOLOGY

1. Végétative axes

Branching in most species of Weinmannia is

from axillary buds, accompanied by the contin-

ued growth of the apical bud, and the architect¬

ure cooforms to Rauh’s rnodel (KELLER 1996),

However, in several species of sect. Leiospermum

the apical bud of growing shoots can be aborted

and branching is then dichotomous. Most of

these species show a mixture of branching patt¬

erns, but in Weinmannia dicbotoma Brongn. &

Gris, branching is almost exclusively dichotq-

mous. Occasionally four buds at a node will

develop, producing four stems of equal domin¬

ance (e.g. W dicbotoma, W dffinis A. Gray). The

buds that develop in this instance are latéral to

the axillary buds, and in a plane perpendicular to

the line: leaf- axil - (apical bud) - axil - leaf, and

are referred to as latéral, auxiliary buds; again,

the apical bud is aborted.

In Weinmannia, as in most other Cunoniaceae,

there is a pair of small, flattish, usually rounded

to ovate, leaf-likc structures betwcen the pétioles

at each node, usually reftrred to as interpetiolar

stipules (Rl'TLSHAUSER &t DlCKISON 1989; Fig.

2A,B; 3Q). MELVILLE (1972) described them as

bud scales and considered that they were not true

stipules but that each was équivalent to an entire

leaf. At végétative shoot apices, they cover the

developing buds and can be either persistent or

caducous as the leaves enlarge; in the later case,

they leave a distinctive annular scar. Similar

structures cover developing (partial) inflores¬

cences and are also found at nodes within the

inflorescence in sect. Leiospermum where the

leaves are wholly or partly suppressed. When

found in the inflorescence, the stipules are usual¬

ly more oblong-elliptic than at végétative nodes.

Stipules can be variable in size and shape on a

single specimen and they are of varying value as

taxonomie characters. They are offen absent from

the fertile shoots of some species and tend to be

larger and more persistent on végétative shoots.

The arrangement of leaves is always opposite

and decussate, giving a distinctive appearance

(Fig. 2A). Compound leaves are usually impari-

pinnate though the terminal leaflet is sometimes

absent or fused to a latéral one. The petiole and

leaf rachis can be winged or not, and sometimes

the distinction berween these two States is not

marked. In Malesian and Pacific species, the

wings are never as well developed as in some

American ones. The size and shape of the leaves

and leaflets, and the amount and distribution of

indumentum, are often taxonomically signifi-

cant. l.eaflet size i.s small to medium, and in

contrast to the Neotropics, no species is micro-

phyllous. The rnargin is almost always distinctly

dentate or crenate, and in W. serrata Brongn. &

Gris, some-what sinuate. In young leaflets the

ends of the teeth are pronounced and probably

glandular.

The venation is rather uniform. Secondary

veins usually start to branch and anastomose

Fio. 2.— A. Weinmannia fratioea, ’oliage ot immature shoot,

showing opposite and decussate oompound leaves and large,

rounded, salvertorm Interpeliolar stipules at nodes; young folia-

ge reddish. ( Hopkins et al 5001a. Ml. Maiang. Sarawak) —B,

Weinmannia clemensiae. loliage and immature fruits.

Intructescence a pair ot dyads with short padundes. Note large,

rounded stipules, ( Hopkins A Bradiord 50W. Pig Hill, Kinabalu,

Sabah.l.— C, D, Weinmannia fraxmea " dulitensîs C, racemes

ot male flowers; D, base ot inflorescence showing 4 dyads sub-

tended by a pair of opposite petiotes, Noie ffowers inserted in

fascicles. ( Hopkins & Bradiord 5014, Pig Hill. Kinabalu,

Sabah).—E, Weinmannia richii, fruits at dehiscence.

Infructescenoes of dyads with very short pedundes. ( Hopkins S

Bradiord 5028. road N from Monosavu Dam. Viti levu. Fiji).— F,

Weinmannia serrata, immature fruits. Infrudescence a central

pentad and one latéral trlad, with one raceme missing.

(Bradiord 632, Col d'Amieu, New Caledonia).—Ail photos by

J.C. Bradford.

ADANSONIA. sér. 3 - 1998 • 20(1)

Weinmannia in western Malesia

ADANSONIA, sér. 3 • 1993 * 20(1)

Weinmannia in western Malesia

before reaching margin but at least part of the

vein terminâtes at the tip of a tooth. Tertiary and

quaternary venation are strongly reticulate.

Leaf/let texture is often coriaceous or at least

chartaceous-coriaceous. Species that grovv at high

altitudes usually have more numerous, smaller,

more coriaceous leaflets than those from lower

élévations. Black dots, which are probably the

bases of caducous hairs, are frequent on the

underside of the leaves in sonie species. Such

leaves are referred to as “puncrate" iti the species

descriptions. Pustules are small, pale protubér¬

ances on both the upper and lower surfaces of

leaflets and they can .tlso occur on other organ.s,

such as the calyx lobes. They do not appear to

have any taxonomie signifîcance nor to be an

artefact of drying (cf, BERNARD! 1964: 162).

Juvénile faîtage and stérile collections.

The characters that are diagnostic of adult foli-

age are rarely évident in juvénile and coppice

shoots. For instance, a sériés of roadside collec¬

tions from stérile individuals 1-3 m high near

Monasavu Dam, central Viti Levu, Fiji (Hopkins

ér Bradford 5027a-h) show a range in variation in

the number, size and shape of leaflets, length of

Fig. 3.—Details of the nodes, buds, flowers and fruits of

Weinmannia A. flower; B. Hower with calyx and corolla remo-

ved to reveal dise lobes, filaments and gynoecium; C, l.s.

through B to show ovules, D, gynoecium with syncarpous ovary

and 2 dise lobes: note groove down style and down side of

ovary: E, calyx lobe with ciliolate margin and 3 glands; F, pelai;

G, anthers, L.H.S, before dahlsoence, R.H S after dehlscence:

H, seed wilh hair tuH at each end and sculptured surlace, note

raphe down midline: J, t.s. ovary (D) to show developing ovules;

K, ouler surface ot one valve of a capsula showlng exocarp

(central pan sirigose) and endocarp exiended as wmgs al both

sides, on réceptacle, the calyx. lobes fallen but dise lobes persis¬

tent L, inner surlace ot K. note persistent centrai cotumn and

minutely sculptured shiny endocarp; M developing Inflores¬

cence, one latéral Iriad shown in axii of petiole. note stipular

scar between pétioles, "collars" arourid latéral peduncles and

conical dilate buds at base ot latéral peduncles (latéral auxiliary

buds): at base of triad there is a stipule (partly lolded baoK) and

a pair ot muen redueed leaves: N, section of axis of a raceme

typical ot secî. Leiospermum, thé pedicels inserred singly ig the

axils of th6 bracts: P. section ot axis ol a raceme in sect.

Fasciculata , the pedicels inserted in tascides In the axils of the

bracts; Q. stipules between leaf petiotes ai most distal riode of

shoot protecting apical bud Scale bars i mm unless otherwise

indicated. |A-F, J, W. patlettëis ( McPherson 3403); G, W. serra-

ta (Bernardi 12816); H, W. serrais ( Balansa 204), K, l, W pait-

ensis (Veiilon 296); M, W. serrata (Bnnon 746); N, W, serrata

(Balansa 2859); P, W. frarinea "dulitensf (Hopkins & Bradions

5014); Q, W. serrata (Lécard s.n., reçu 20 Oct. 1879)].—Drawn

by H.C.F. Hopkins.

intemodes, vestiturc, shape ot rachis, shape and

persistence of stipules. Both W. richii and W.

affims occur in this area but the stérile collections

were rarely consistent with the adule foliage of

either of them, Sonie of this variation could per-

haps bc- due to hybridization and the coloniz-

ation of disturbed roadsides by hybrid swarms.

Characters of mature foliage are best seen in

leaves at the periphery of the crown m mature

individuals (i.e. foliage from just below the

inflorescences).

2. Structure of the inflorescence

{with J.C. Bradford)

The parts of the inflorescence are organised hier-

archically, and in Weinmannia, there are rypically

three levels of organisation; 1) flowers, borne

along an unbranched axis, here termed a raceme;

2) racemes, organised into metameric units, each

referred to here as a partial inflorescence (PI); 3)

parmi inflorescences, borne along the main

stem Togerher, the structure of a raceme, the

way the racemes are organised into a PI, and the

arrangement of the Pis along the main stem,

constirute the inflorescence. Although there is

sorne interaction amongst levels of organisation,

with higher levels influencing lower ones, most

variation within one level of organisation appears

to be independent of variation at another level.

For this reason, each level of organisation will be

discussed separately

There are several terms that require short défin¬

itions so that the descriptions of the inflores¬

cences and their component parts can be made

clearly and coneisely. A bract is a minute, often

carinatc blade along the axis of a raceme and it

subtends a flower or a group of flowers. A bud is

an undeveloped shoot or flower, which in

Weinmannia is normally cnclosed by a bract (flow¬

ers) or a pair of stipules (shoots). A stipule is a

small leal-like structure at a node, decussate with

the crue leaves that may themselves be partially

ot wholly suppressed at nodes within die inflor¬

escence. A metamer is a unir which consists of an

internode, the node at its distal end, and the

organs and meristems (buds) associated with that

node. The term “main stem” is used for any

dominant, linear axis of growth.

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

a) Racemes .—The flowers of the Malesian-

Pacifîc species are pedicellate and borne in

racemes ca. 5-12 cm long. The bracts usually fall

when the flowers are still at the bud stage but

occasionally they persisr until fruiring. Afrer they

hâve fallen, a minute scar is usually visible. The

individual flowers or Fascicles are rather irregular-

ly arranged on the axis of each raceme and insert¬

ion is never truly verticillate though it may

superficially appear to bc. Racemes may bear

flowers for the wbole of their length or usually

they are devoid of flowers towards the base. The

flowers in orie raceme open more or less simultan-

eously.

— Sect. Fasciculata, Several pedicels are insert-

ed in a small cluster in the axil of a bract (Fig.

3P). Occasionally flowers are inserted in fascicles

for the most part but wirh some inserted singly

towards the apex of the raceme, and in a few .spe¬

cies, the fascicles rend to dissociate from the

bracts during development. BtRNARDl (1964:

132) used the term ‘pseudoraceme in reference

to sect. Fasciculata, but since this term is rather

ambiguous, it is not used hcre.

— Sect. Leiospermum. The pedicels of the flowers

are inserted singly in the axil of a bract (Fig. 3N).

b) Partial inflorescences .—In general, a PI is

any subunit(s) into which the inflorescence can

be divided. As used here, it consists of racemes

and one or more stérile stem sections which torm

a developmentally integrated unit whose structure

is predictable wichin different taxonomie groups.

However, sometimes (though rarely in this

région) the inflorescence of Weinmannia is com-

posed of individual racemes inserted in leaf axils,

in which case the racemes could be considered to

be the Pis.

— Sect. Fasciculata. A partial inflorescence is

limited to a single metamer. An even number of

racemes (2 or 4) are inserted at lhe distal node of

a short, stérile stem segment (referred to as a

peduncle in the species descriptions) to form

either a dyad (Fig. 4A) or a tetrad (Fig. 4B). Leaf

production is usually totally supptessed at the

node from which the racemes arise (exceptions

seen in W. pullet)

There is typically a small bud in the angle bet-

ween the central pair of racemes at the apex of the

peduncle. This bud has the potential to continue

végétative growth after flowering. LJsually it is

dormant during flowering (prolcptic) or rarely

sylleptic (i.e. no cessation of growth).

— Sect. Leiopsermum A partial inflorescence

can consist of more than one metamer. Usually,

an odd number of racemes ate inserted at one or

more nodes along the axis of the PI as the termin¬

al meri.stem develops into a raceme. Theretore,

when the PI consists of one metamer, there are 3

racemes (callcd a tri ad, Fig. 4F), when of two

metamers, there are 5 racemes (a pentad , Fig.

4G), and when of three metamers, there are 7

racemes (a beptad). In a few species the Pis are

dyad.s (Fig. 4H) and the bud between the bases

of the racemes has aborted (i.e, fai lcd to develop

into a third raceme; exceptions in New Zeatand

are deseribed below). The basal intemode of a PI

may be loosely termed the peduncle.

Leaf production can be either partially or total¬

ly suppressed within each PI and when leaves are

produced, they are much reduced (e.g. Fig. 4J,

level Y). There are often stipules at nodes within

a PL Quire often there are “collars” around the

base of the pcduncles of latéral Pis (Fig. 3M).

Thèse appear to be the rem riants ot a pair of stip¬

ules which once covered the developing PI and

were fused to each other at the base. They are

not usually seen around the peduncle of the

central PI in sect. Leiospermum, nor around the

pedundes in sect. Fasciculata.

c) Inflorescences .—An inflorescence is compos-

ed ol one or more partial inflorescences. Usually

ail the flowers in one inflorescence open more or

less simulraneously.

— Sect. Fasciculata (Fig 4A-E). The Pis devel¬

op in opposite pairs from latéral meristems at a

node on the main stem, ce. they are usually in

the axils of leaves. Each PI is a dyad or a tetrad.

The most distal végétative node on a stem usual¬

ly produces more pairs of Pis than do subdistal

nodes (see discussion of supernumerary buds

below) and often the initiation of Pis is confined

to the most distal node on the main stem.

Since the Pis are latéral, the apical meristem of

the main stetn remains indeterminate, It is

normally dormant during flowering and contin¬

ues végétative growth after fruiting (prolepsis) or

rarely it may be sylleptic. In exccptional cases,

the apical meristem gives rise to a media! PI but

ADANSONIA. sér. 3 • 1998 • 20(1)

Weinmannia in western Malesia

Fig. 4.—Inflorescence structure in Weinmannia. A-£ sect. Fasciculata, A, B: partial inflorescences: A, dyacf B. tetrad: C. inflores¬

cence typlcal of W fraxînsa group, composed of 4 dyads and a pair of latéral supernumerary buds at the rnost distal végétative node

and single dyads in the axils of a pair of more proximal leaves; apical bud proleplic: D. inflorescence typical of W. urdanetensis

group, composed of a pair of axillary dyads: apical bud proleptic: E. as D, but apical bud sylleptic.. F-L, sect, Leiospermum F-H:

partial inflorescences - F tnad; G penlad; H, dyadwrth central raceme aborted; J, inflorescence typical of W. parvitlora. consisling of

a heptad wirh 2 additional pairs of racemes inserted af node Z. partially suppressed leaves at node Y; K, inflorescence typical of W.

dicholoma, with 2 axillaty partial inflorescences (both dyads here) and apical bud aborted. Note prominant auxiliary buds In latéral

positions at base of pedundes; L, inflorescence of 3 partial inflorescences (ail triadsf.A: bud: ®: peduncle; x: raceme.—Drawn by

C. Monnie.

ADANSONIA. sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

végétative growth cari agairt continue from the

indeterminate apical merisreni within tKat PI.

In most species (e.g. W. fmxinea , W. hutchinson-

ii Merr.) there is typically more than one bud per

axil with the porenrial to produce a PI and these

“supernumerary buds” are insetted in sériés.

One, two or three buds raay develop per axil,

producing one to three pairs of Pis. Level X of

Fig. 4C shows 2 pairs of dyads and one pair of

dormant supernumerary buds. The racemes are

arranged in a plane that is perpendicular to that

formed by the line: leal - (bud) - peduncle -

peduncle - apical bud - peduncle - peduncle -

(bud) - leaf (see Fig. 2D).

In another group of species (e.g. W.

urdanetensis, W. pullri) there are typically only

two Pis (here dyads) which are inserted in the

opposite .axils of the most distal pair of leaves

(Fig. 4D). Sometimes there are additional Pis in

the axils of more proximal leaves as well but

there are no additional buds in sériés at rhe same

nodes as the dyads (i.e. there is only one bud per

leaf axil with rhe porenrial to produce a PI). The

apical bud of the main stem can be either prolep-

tic or sylleptic (Fig. 4E), in which case the dyads

that were initially at the most distal node become

subdistal.

— Sect. Leiospermurn (Fig. 4F-L, 5). Partial

inflorescences are usually positioned medially

and often laterally too. In some cases, the apical

bud of the main stem aborts and only latéral Pis

are produced. The switch to flowering usually

terminâtes the végétative growth of the main

stem. In any one inflorescence rhere is usually an

odd number of Pis (usually 1 or 3; rarely 2, a cha-

racter State seen most Irequenrly in some species in

New Caledonia); when there is more than one,

they are not always identical and the media] PI

often contains more metamers rhan the latéral

ones.

In the simplest examples (e.g. Weinmanniti mar-

quesana F. Br.) the inflorescence consists of a

single PI (a triad or pentad) which develops Irom

the apical bud of the main stem; the lower pair of

racemes may be in the axils of citlier hilly develop-

ed, partially suppressed or rotally suppressed

leaves. When an inflorescence consists of more

than one PI, there are usually 3 rriads (Fig. 4L.) or

a central pentad with a pair of axillary triads (as

often seen in W serrata and W ajjinis), and occa-

sionallv more complex arrangements are found.

A unique structure is seen in Weirtmannia parvi-

flora (Fig. 4J). The inflorescence illustrated

consists of a central heptad with a pair of partial¬

ly suppressed leaves at one node, and 2 addition¬

al pairs of racemes at the lowest reproductive

node. These additional racemes irise in a plane

perpendicular to that formed by the line: leaf -

raceme - central axis - raceme - leaf (see Fig. 4J,

level Z), i.e. they are latéral to the axillary raceme

and not inserted in sériés. One possible explanat-

ion for their presence is that the three racemes

arising on either side of this node represent a pair

ot reduccd triads in which the peduncle has not

developed. Evidence for this is the occasional

growth of a tiny section of peduncle in this axil

Fig. 5.—Inflorescence structure in the New Zealand species of Weinmannia. A. B, W. sylvicola. A, 2 sequential pairs of dyads

(pentad minus central raceme?): apical bud aborted; B, as A with an additional partial inflorescence (dyad) in an axillary position at

subdistal node; C, D, IV. racemosa C. inflorescence a médian dyad. apical bud proleptic; D. as C but apical bud sylleptic, and con-

tinuing to produce dyads and leaves at sequential nodes.—Drawn by H.C.F. Hopkins.

ADANSONIA. sér. 3 • 1998 • 20(1)

Weinmannia in western Malesia

(see illustration of W. rarotongensis Hemsl., Fig.

7G in Hopkins & Florence 1998).

In three of the New Caledonian species and

occasionally in other Pacific ones, the apical bud

of the main stem may abort. In Weinmannia

dichotoma for instance, the inflorescence fre-

quently consists of two latéral Pis that develop at

the most distal leaf-bearing node (Fig. 4K).

These Pis can be either triads (Fig. 4F) or dyads

(Fig. 4H,K), where the apical meristem within

the PI has aborted also. At the node from which

the Pis arise, there are often promïnent latéral

buds in a plane perpendicular to that of the

leaves (latéral, auxiliary buds, Fig. 4K). They are

most obvious in W. dichotoma but occur in other

species as well.

In both species from New Zealand,

Weinmannia sylvicola Sol. ex A. Cunn. and W.

racemosa, the inflorescence structure is unique

although ît can still be understood as part of the

pattern fourni in sect. Leiospermum (Fig. 5).

In Weinmannia sylvicola, rhe inflorescence is

similar to those already described for sect.

Leiospermum in which Pis consist of more than

one metamer and they can develop in both

médial and latéral positions. However, W. sylvicola

is distinctive in that the apical meristem usually

aborts (rarcly not, e.g. Colenso s.n. where it

appears dormant), so that the PI in a médial

position resembles a pentad without the central

raceme (Fig. 5A). In rhis instance, the apical

meristem of both the PI and the main stem is the

same. Apical meristems commonly abort in three

of the species from New Caledonia but then the

PI consists of only a single metamer. though in

some other Pacific species where the apical meri¬

stem of the main stem somerimes aborts, latéral

Pis may be pentads or heptads (e.g. W. purpurea

L.M. Perry) and consist of more than one meta¬

mer. As in orher pentad inflorescences, leaf pro¬

duction in W. sylvicola may be normal, partially

or totally suppressed at the node front which the

lower pair of racetnes arise. When latéral Pis

develop, they usually consist of only one meta¬

mer and often they develop at only one side to

produce an asymmetrical inflorescence (Fig. 5B).

Superficially, the inflorescence of Weinmannia

racemosa appears to hâve little in common with

those of other members of the section. However,

its homology can be deduced if some assump-

tions are made abour its evolutionary relation-

ships. Here, we suggesr that W. racemosa is most

closely related to W. sylvicola (the only other New

Zealand species), and that overlapping forms of

these two species can be identified and used to

link the unusual inflorescences in W racemosa

with those rypical of sect Leiospermum.

In Weinmannia racemosa there is always a single

PI in a médial position and it can consist of

either one (Fig. 5C) or two meramers. When two

metamers develop the structure appears similar

to a common variant of IV.' sylvicola in which

there is no latéral PI (Fig. 5A), excepr that in rhis

case, the apical bud is not aborted. There is thus

a continuity in fbrm berween W. racemosa and W

sylvicola, and between W. sylvicola and other

Leiosperma. The fate of rhe apical bud in W.

racemosa makes it unique within sect.

Leiospermum in that végétative gruwth can contin¬

ue from within the inflorescence. This, together

with the absence of latéral Pis and the generatly

lesser number ot metamers in any PI, is the rea-

son for the unusual appearance of the inflores¬

cence in W. racemosa. The apical bud (of both

the PI and the main stem in this case) can be

either dormant or commonly not, when it goes

on to produce further leaves and Pis (Fig. 5D).

3. Flowers

While basic floral structure is extremely uni-

form among different species (see Fig. 3A-G),

flowers can be morphologically bisexual (with

filaments approximately the same length as the

gynoecium so that anthers and stigmas are at

about the same level, Fig. 6C), male (with long

filamenrs, the anthers far exserred beyoncl the

stigmas, the ovary small and the styles tiny and

incurved, Fig. 6B) or female (with short fila¬

ments, the ovary often large and the styles far

exceeding the stamens, ejther straight or çup/jng

outwards, Fig. 6A). However, there is often a

considérable range in the length of the filaments

and occasionally they are not clearly long or

short, and then the gender of the flowers is equiv-

ocal. Usually the anthers are of similar size in

male and female flowers, though unusually small

anthers are found, for example, in the apparently

ADANSONIA. sér. 3 ■ 1998 • 20(1)

Hopkins H.C.F.

Fig. 6.—Androecium and gynoecium from morphologically female (A), male (B) and bisexual (C) flowers of Weinmannia .—Drawn by

J. Lemeux.

bisexual flowers of W. fraxinea from the

Solomons.

There is only preliminary information on the

functional poteniiaJ of flowers of different mor-

phological types. In W. denhamii Seent. from

Vanuatu, the anthers of botli male and female

flowers contain pollen, although in the female

flowers it appears to be less well formed (R.D.

Hoocland in lut. ro J. Florence. 19 May

1987). In W. parviflora from Tahiti, no pollen

was found in the anthers of morphologically

female flowers {Florence 9089, 9091), while the

anthers of morphologically males ones ( Florence

9089 , 9090) did contain pollen (R.D.

HOOGLAND unpublished notes).

Sexual expression and breeding System. In

many species, ail the flowers on one herbarium

specimen, which mighr contain several inflores¬

cences each composed of several racemes, are of

the same morphologica! type and there is no évi¬

dence to suggest rhat some species are not either

simply djoecious (e.g. W hutchimonii , W. luzon-

iensis S. Vidal, W aphanoneura Airy Shaw) or

hermaphrodific (e.g. W. serrata and other species

from New Caledonia, W. racemosa and W. sylvic-

ola from New Zealand, W furfuracea H.C.

Hopkins and W. descombesianA from Sulawesi).

The expression of sex is more complex in some

other species. For instance, a number are poly-

gamodioecious (S MIT H 1952), with male and

female flowers on separate plants and some

bisexual flowers, usually on staminate plants in

this case (e.g. numerous species front small

Pacific islands). In these species, bisexual flowers

are usually less common tban unisexual ones,

and they may either be mixed in the same inflor¬

escence as the male flowers or in separate but

contemporaneous inflorescences. In some species

(e.g. IV marquesana, W. denhamii, W. rapensis)

there are also temporal changes in gender, with

cycles of different sorts of flowers on one plant,

indicated by the presence of male flowers at

anrhesis and mature fruits (usually from bisexual

flowers) in separate inflorescences. Temporally

labile species may contain some stable indivi-

duals and others that change gender within and

berween seasons.

In some predominantly dioecious species (e.g.

Weinmannia urdanetensis) a few individuals

appear to be tnonoecious. The ntost complex

pattern is seen in W fraxinea where the sexual

System varies geographicatly and some popula¬

tions may perhaps be apomictic. Betrcr descrip¬

tion of the more complex sexual Systems clearly

requires detailed, long-term fteld studies.

Compatibility mechanisms hâve not been stud-

ied. There is circumstantial evidence of inrerspe-

cific hybridization, e.g. in central Viti Levu,

where small, infertile roadside plants showed a

wide range of végétative characters, and were

likely to hâve been hybrids between W. richii

(sect. Fasciculata) and W. affinis (sect.

Leiopscrmurn) (HOPKINS &( BRADFORD pers.

obs.).

Floral biology andpollination, The flowers of

Weinmannia are massed together in fairly dense,

spike-like inflorescences that are arranged around

the periphery of the crown, projecting upwards

ADANSONIA. sér. 3 • 1998 • 20(1)

Weinmannia in western Malesia

(Fig. 2C). Individual Flowers are bowl-shaped

with the anthers and stigmas readily accessible to

potential pollen vectors that land on die racemes

(i.e. non-restrictive). The Flowers are usually

white or créant, sometimcs pink, and described

as scented or not. Nectar is produced iti

Weinmannia racernosa (Thompson cited in

WARDLE 1966) althongh none was seen in W.

fraxinea or W. affinis (Hopkins & Braoford

pers. obs.). The dise lobes are pale and inconspic-

uous in W. fraxinea in Bornéo while in W. affinis

and W. vitiensis in Fiji, they are darlc and contrast

strongly with the pale corolla, There hâve been

no detailed studies oF pollination but becs are

probably the main pollen vectors. The Flowers ol

W. racemnsa in New Zealand are visited by

insects including becs, and by the Farsonbird or

“tui”, Prosthernadera novaeseelandiae (Wardle

1966). Racemes of Weinmannia also attract a

variety oF (lies and beetles (pers. obs.). A high

proportion oF female or bisexual Flowers on any

raceme produces Fruit.

4. Fruits and seeds

The capsules (Fig. 2E; 3K,L) are remarkably uni-

form, with only minor variation between spccies

in size and indumentum. Variation in the per-

sistence oF the calyx lobes bas been used as a taxon¬

omie character but is variable in a few species. In

sect. Leiospermum , thete is a well developed per¬

sistent central colunui (GoilLEY 1983; Fig. 3L)

from which the latéral wings of the endocarp

separate on dchisccncc. In sect. Fasciculata, this

column is often moie weaklv developed. or short-

er or absent, when the tissuc chat otherwise Forms

the column remains attached to the endocarpic

fringes of the valves. The capsules are usually

referred to as septicidal, but the presence of a free

central column after dehiscence means that they

are septifragal (see SPJUT 1994).

The seeds are small and che surface of the testa

has a complot pattetn of sculpturing (DlCKISON

1984; Werb & Simpson 1991; Fig. 3H). In most

species there Ls a tuft of comose hairs at each end.

although in a few, there are short hairs wtdely dis-

tributed over the entire surface. There is some

variation in che length and texture of the comose

hairs which appears to affect the length of time

the seeds are retained wichin the capsule after

dehiscence, and this may affect their dispersability.

Dispersai. The seeds are probably wind-dispers-

cd (e.g. Wardle 1966; Hopkins & Bradford

pets. obs. in Fiji) although exozoochory has been

suggested (GUPPY in FLORENCE 1982: 32, for

Weinmannia parviflora), the hairs oit the seeds

perhaps sticking to birds' feathers. Long-distance

dispersai must be reasonably effective since the

genus has rcached many volcanic islands in the

South Pacific.

5. Field characters

Life form is rather plastic in some species which

can bc trees, shrubs, dwarf shrubs and occasion-

ally woody epiphytes, depending upon che végét¬

ation type, altitude and exposure. Bark Ls often

variable within species, and rarely distinctive.

Tnner bark is often pinkish, brown or reddish

brown. Sap wood is white, straw, pinkish, some¬

times orange-brown, somerimes rurning purple;

medium ro hard. Heart wood is pink-cream to

red ot yellow-brown. Young stems and leaves are

often conspicuously red (Fig. 2A), mature leaves

often darkish green above, shiny; old leaves

sometimes rurning bright red. Inflorescence axes

are sometimes reddish. Flower buds are light

green to pink or red. Flowers are creamy white to

pink (Fig. 2C,D), rarely light green or purple (W

purpurea), often with a slight swect fragrance; the

calyx is pale green, yellowish green or reddish;

the corolla and filaments are usually white; the

anthers pinkish, cream or straw; the dise yellow-

orange or dark red-purple; the ovary light green

or pinkish; the styles are pink at the base and the

stigmas are white to brown or purplish. Young

fruits are greenish, pink or reddish (Fig. 2B,F),

turning crimson or rarely purplish green at

maturity; dehisced capsules are brown (Fig. 2E).

WEINMANNIA L.

Syst. Nat., cd. 10, 2; 997, 1005, 1367 (1759), nom.

cons.; Juss.j Gen. PI.: 309 ( 1789); Lam., Tabl. Encycl.

I 2, t. 313 (1793); Willd., Sp. PI., ed. 4, 2: 296, 436

(1799); Benth. & Hook. f„ Gen. PI. 1: 653 (1865);

Engl., Linnaea 36: 592 (1870); Engl., in C. Mart., Fl.

Bras. 14 (2): 156 (1871); Engl., Nat. Pflanzenfam.,

ADANSONIA. sér. 3 - 1998 • 20(1)

Hopkins H.C.F.

111, 2a: 101 (1891), ed. 2, ISa: 250 (1930): Bail!.,

Hist. PI 3; 373, 447 (1871): Bader, Nova Acta Leop.

23 (148): 170, 430 (1960); Bcrnardi, Candollea 17:

123 (1961), 18: 285 (1963); Bcrnardi, Adansonia, ser.

2, 3: 404 (1963); Bcrnardi Bot. (ahrb. Syst. 83: 126,

185 (1964); Bcrnardi. in Humbert, Fl. Madag., Cun.

93: 3 (1965); Hutch., Gen, Fl. PI. Dicot. 2: 9 (1967).

Windmannia P, Browne, Giv. Nat. Hist. Jamaica: 212

(1756), nom rejic.; Kuntze, Revis. Gen. PI. 1: 228

(1891).

Arnoldiii Blunic, Bi/dr. Fl. Ned. fnd.: 868 (1826),

non Cass. (1824).

Leiospermum D. Don, Edinburgh New Philos. J. 9: 91

(1830).

Pterophylla D. Don, Edinburgh New Philos. J. 9: 93

(1830).

Ornithropbus Bojer ex Engl., Linnaea 36: 636 (1870),

pro syn.

Type. — Weinmannia pinnata L. from Central and

South America and the Antilles.

Generic description of Weinmannia

in Malesia and tlhe Pacific

Trees or shrubs, rarely epiphytes and perhaps

rarely stranglers. Branching sometimes dichot-

omous in sonie species, usually so in a minority

of species; shoots sometimes slightly thickened or

flattened at nodes, internodes terete, the bark

ftnely grooved; pale lenticels ofren promurent.

Indumentum of simple hairs, variable in amount

and distribution. Interpetiolar stipules 2 per

node, elliptical, ovate, orbicular to subreniform,

often with minute adpressed liaits, apex rounded

or pointed, margin entire or roothed, a pair of

opposite stipules often salverform in juvénile

foliage, sometimes caducous in adult foliage, the

bases leaving a prominent annular scar around

stem at each node aller abscission. Colleters

purpie-red, minute (< 0.1 mm long), frequent at

nodes and visible afrer stipules have fallen, non-

functional? Axillary and apical buds covered by a

pair of velutinous or sericeous stipules (bud

scales). l.eaves opposite and decussate, simple

(unifoliolace?), trifoliolate or imparipinnate (rare¬

ly paripinnate), usually petiolate; petiole and

rachis terete, caniculate, or semiterere and flatten¬

ed on adaxial side, sometimes narrowly winged,

especially distally towards the point of insertion

of each pair ol leallets; leaflets chartaceous to

coriaceous; latéral leaflets often asymmetrical at

base, often smaller proximally in a given leaf;

margin tooched or crenulate, the crcnations acro-

scopic, margin usually minutcly thickened and

minutely revolutej seoûndary veins usually arcu-

ate, branching belore reaching margin, one bran¬

di arching towards apex and the other towards

base to join with tertiary veins arising from a

more proximal secondary vein; blade sometimes

hlack-dotted below, sometimes pustulate.

Inflorescence composed of racemes, usually

arranged in groups on a stérile axis on wliich leaf

development ïs partially or wholly suppressed, to

form a developmentally integrated unit, rarely

not part of such a unit and arising directly from

axillary buds on the main leaf-bearing stem; part¬

ial inflorescences either axillary or terminal or a

combination. Racemes up to 12 cm long with up

to 100 or more flowers, the basal part usually

bare; axis of racctne longitudinally ridged and

often minutely hairy (hairs < 0 1 mm long),

sometimes densely so. Floral buds inserted on

the axis of the racemes either singly or in small

fascicles, each bud or fascicle subtended by a

bract; bract often carinate, often histute on

abaxial surface, often caducous, Flowers small,

unisexual or bisexual, hypogynous or somewhat

perigynous, perianth and androccium 4-merous

or rarely a fcw flowers per raceme 5-mcrous

(commonly 5-merous in sonie areas outside this

région); pedicel usually fllifbrm, often minutely

hairy (hairs < 0.1 mm long); calyx ol 4 lobes,

imbricatc in bud, ± triangular, attached at their

base to the hypanthium at the point ol their

greatest width, apex broadly acutc or somewhat

rounded, glahrous or hirsute on outer surface,

slightly fleshy; corolla of 4 free pctals, not cadu¬

cous (in rhis région), usually elliptical, ovate or

obovate, constricted at the base (attachaient 0.1-

0.2 mm across), apex rounded or irregularly

emarginate, membranous, sometimes minutely

ciliolate; androccium ot 8 sramens, obdiplo-

stemonous, the filaments filiform, inserted in a

single whorJ but alternate ones elongating asyn-

chronously; atithers apiculate, versatile, dehiscing

longitudinally, the valves of each thccum un-

equal; dise of 8 free lobes, alternating with the

filaments, usually ± oblong, obeuneate or slightly

claviform, sometimes broadly obiong with

(langes on either side so that lobes almost form a

ADANSONIA, sér. 3 • 1998 ■ 20(1)

Weinmannia in western Malesia

continuons ring of rissue, glabrous; gynoecium

of 2 (rarely 3) carpels fused at the level of the

ovary; ovary ovoid, glabrous to densely hirsute,

the hairs drying white; locules 2(-3); styles 2(-3),

free, awl-shaped, glabrous exccpr occasionaily at

the base, conduplicate and furrowed on adaxial

side; stigmas terminal, small or sometimes capit-

ate, papillose; ovules ca. 8-16 per locule, placen¬

tation axile.

Fruit small (< 6 mm long), a septicidal or usual-

ly septifragal capsule dehiscing from the apex

into 2 parts which remain attachetl to the récep¬

tacle at their base to fornt a cup-shaped structure

with the styles usually persistent at rhe apex; the

valves composed of 2 layers: exocarp dark brown,

pubescent, strigose or glabrous, smooth or

minutely ridged; endocarp yellow, smooth. with

minute transverse sculpturing on concave surl¬

ace, usually completely adnate to exocarp or

sometimes partially free especially towards the

apex, the septum separaring rhe locules extend-

ing as wing-like projections on either side of the

exocarp at dehtscence; free central column form-

ed from placenta oftc-n présent and sometimes

persisting after the valves hâve fallen; réceptacle

sometimes saucer-like at base; calyx lobes persis¬

tent or not; corolla and dise lobes occasionaily

persistent, Seeds small (< 1 mm long), elliptical in

outline and circtilar in transverse section, generally

2 X as long as broad when fully mature, the surf¬

ace minutely sculptured, raphe sometimes visible

as a minute ridge, usually comose with straw-like

hairs in a ttift at each end, sometimes hairs arising

± ail over the surface, longer at the ends.

About 40 species in Malesia and the Pacific,

belonging to rvvo sections.

1. Sect. Fasciculata Bernardi ex Hoogland &

H.C. Hopkins, sect. nov.

Sect. Fasciulatae Bernardi, Bot. Jahrb. Syst. 83: 132,

158 (1964), nom. inval., sine typus.

TYPE (here designated). — Weinmannia fraxinea (D.

Don) Miq.

Flowers pedicellate and inserted on the axes of

the racemes emirely or largely in fascicles, i.e.

several pedicels arising in a group, the group sub-

tended by a single small bract (Fig. 3P), rarely

inserted singly ( W. descambes'utnd). Inflorescence

composed of 1-3 opposite pairs of partial inflor¬

escences inserted in séries in the axils of the most

distal pairs of leaves; each partial inflorescence

consisting of a single metamet, which comprises

a stérile peduncle and 2 or 4 racemes inserted at

its distal end (to form a “dyad" or a “tetrad", see

Fig. 4A,B); leaves at nodes from which racemes

arise usually totally suppressed; stipules not

usually présent at nodes within partial inflores¬

cences; ‘^0113^ around peduncles of partial

inflorescences absent; bud within cach partial

inflorescence, at apex ol peduncle between the

central raccmcs, présent and usually dormant

during Howering; apical bud of main stem, bet¬

ween central peduncles, présent and usually dor¬

mant during flowering; buds latéral to axillary

buds at most distal leat-bcaring node (latéral

auxiliary buds) absent

Stipules ofien ± orbictüar. Branching not dichot-

omous. Leaf blades not punctate below.

Capsules ± smooth and not ridged; indunient-

um on valves soffly pubescent; after dehiscence,

central column weakly developed; calyx lobes

often but not always persistent in fruit.

20 species in Malesia and the western Pacific,

extending as far east as Fiji.

2. Sect. Leiospermum (D. Don) Engl., Nat.

Pflanzenfam. III 2a: 101 (1891), ed. 2, 18a: 256

(1930).

Sect. Racemosae Bernardi, Bot. Jahrb. Syst. 83: 132,

185 (1964), nom. superfl. et sine typus.

Type. — Weinmannia racemosa L. f.

Flowers pedicellate and inserted on the axes of

the racemes singly, i.c. cach pcdicel arising in the

axil of a small bract (Fig. 3N). Inflorescence

composed l-.3(-5) partial inflorescences, usually

inserted mcdially and lateraUy; latéral Pis usually

consisting ol one metamer [dyad (Fig. 4H,K) or

triad (Fig. 4F,L)]; sometimes lacking the basal

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

stérile stem section in W pannflord)\ médial Pis

consisring of one metamer (dyad or triad), two

metamers [tetrad (Fig. 5A) or pentad (Fig. 4G)]

or three metamers (heptad); apical bud of shoot

usually fertile (developing into a raceme in

triads, pentads and heptads), sometimes aborted

(in dyads and tetrads) or rarely dormant or végét¬

ative (e.g. W. racemosa, Fig. 5C,D); leaves at

nodes from vvhich racemes arise not suppressed,

partially suppressed or totally suppressed; stipules

sometimes présent ar nodes within partial inflor¬

escences: “collars” around peduncles of latéral

partial inflorescences often présent (Fig. 3M);

apical buds within each latéral PI either develop-

ed into a médial raceme or aborted; latéral auxil-

iary buds (i.e. laverai to axillary buds) présent at

most distal leaf-bearing node and often at other

nodes also.

Stipules olten ligulate, elliptical or ovate, the

apex rounded or pointed. Branching sometimes

dichotomous. Leaf blades often punctate below.

Capsules often minutely ridged; indumentum

on valves strigose or absent; afrer dehiscence,

central column usually well developed and per¬

sistent; calyx lobes often falling in fruit.

There are 17-19 species in the Pacific, extend-

ing westwards to Papua New Guinea (Bismarck

Archipelago and Karkar Island).

Notes on the species accounts

1. Végétative characters. Because of variability in

the leaves depending their position, even within

the crown of one tree, and on the âge of the

plant, only leaves front fertile shoots are describ-

ed. Dimensions given are fox the largest leaves

per specimen. Characters for juvénile or coppice

shoots are given separately where known.

Terminology for the indumentum follows

Hewson (1988)

2. Inflorescence. The most frequent structures

are described but nor ail variants.

3. Flowers. The insertion of the pedicels on the

axes of the racemes is scored in bud where pos¬

sible as this is sometimes less dear after the flow-

ers hâve opened and the bracts hâve fallen. The

length of pedicels is given Ibr flowers at anthesis

and may be slightly longer in the fruiting stage.

The length of the calyx lobes is measured from

rheir point of attachment to the réceptacle to

their apex (Fig. 3E). The length of the style

includes the stigma.

4. Breeding System. Species are described as her-

mapbroditic, dioecious or polygamodioecious

bascd on the morphology of the flowers (Fig. 6).

The reproductive potcntial ot flowers of different

sorts lias notbeen investigated.

5. Ficld characters. Many field characters are

common across the genus and are not always

repeared in each species account.

6. Types. In species accounts for which HCFF1 is

rhe sole aurhor, ! is used to indicate type spécim¬

ens that hâve been studied. Collection details for

types are not usually repeated in the list o( mate-

rial examined.

7- Specimen citations. For those species where I

bave seen relatively few collections, ail material

examined is listed. For those where lhere is much

more material. a sélection is cired to illustratc the

morphological and geographical range of the spe¬

cies.

8. Abbreviations used in citations of specimens.

Gn. - Gunung (mountain), Bt. = Bukit (hill),

Mt. - mountain, Isis. = island, Kamp. = kam-

pong (village), R. = river, Rd. = road, Sg. =

Sungai (stream or small river); juv. = juvénile, fl.

- flowers, fr. - fruits, yfr. = young fruits, st. = sté¬

rile; N = north, S = south, C = central, nr. = near,

ft. = feet, Distr. = Distria, Prov. = Province, Div.

= Division, Ond. afd. = Onderafdeling (subdivi¬

sion).

Five species are recognised from western

Maiesia, the Lesser Sunda lslands and the

Moluccas, ail belonging to sect. Fasciculata.

Weinmannia fraxinea is widespread and variable,

and considerably more than half the collections

from Maiesia belong to this species. Weinmannia

clemenisae is endemic to stunted foresr on ultra-

mafic soils on Mt. Kinabalu in Bornéo, W.

aphanoneura occurs in Bornéo and Sumatra, and

W hooglandii is from high altitude in Peninsular

Malaysia. Weinmannia furjuracea is known from

this région by a single specimen from Seram

(Rutten 2231) and is described with the other

species which occur in Sulawesi, in Part 2

(Hopkins 1998a).

ADANSONIA, sér. 3 • 1998 • 20 (1)

Weinmannia in western Malesia

Key to the species of the Malay Peninsula, Greater and Lesser Sunda Islands and Moluccas

1. Indumentum on young stems, leaf rachises, inflorescence axes and lower surface of leaflets red-brown, often

scurfy; apex of latéral leaflets obtuse (Seram).5. W. furfuracea (sec p. 49)

1’. Indumentum on young stems, leaf rachises, inflorescence axes and lower surface of leaflets grey to golden

brown, puberulerit to tomentose, not scurfy; apex of latéral leaflets acuminate, acute or obtuse.2

2. Latéral leaflets small. 1.4-2.7 * 0.5-1 cm, boat-shaped or t oblong; young shoots tomentose-velutlnous .... 3

2’. Latéral leaflets larger (2.2-)4.2-8.5(-12) x (0.8-)1.2-3,5(-4.5) cm, Janccolate, clliptical, narrowly obovate to

ovate; young shoots t glabrous, puberulent or tomentose ........4

3. Leaflets bulîate and the margin strongly recurved so Icaflct is boat-shaped; latéral leaflets 6-13 pairs per leaf

(Mt. Kinabalu, Sabah) .. .. . .2. W. clemensiae

3’. Leaflets fiat, not bullate and the margin flat or roinuteJy revolute; latéral leaflets' 4-S pairs per leaf

(Pen'msular Malaysia) .4. W. hooglandii

4. Latéral leaflets narrowly elliptical to narrowly obovate, the base cuneatc, the apex acute to obtuse, drying

chesrnut-brown, 1-4 pairs per leaf; leaf rachis sometïmcs narrowly winged; inflorescence of 1-2 pairs of

dyads . 3. W. aphanoneura

4’. Latéral leaflets, lanceolatc, clliptical or ovate, not obovate, the base usually unequa) and varying front cune-

ate to rounded, rhe apex narrowly acute to acuminate, usually drying grcenish-brown, 1-8 pairs pçr leaf; leaf

rachis terete, not winged; inflorescence of 1-3 pairs of dyads or terrads ...1. W. fraxinea

1. Weinmannia fraxinea (D. Don) Miq.

Fl. Ned. Jnd. 1(1): 718 (1856); JSm. ex D. Don,

Edinburgh New. Philos. J. 9: 93 (1830) pro syn.];

Engl., Linnaea 36: 644 (1870), Nat. Pflanzenfam., ed.

2, 18a: 256 (1930): SchefE, Nat. Tijdschr, Ned. Ind.

Batavia 34: 40 (1874); Pamp., Ann. Bot. (Rome) 2;

89 (1905); Mcrt , Inicrpr. Hcrb. Amboin.: 244

(1917); Bernardi. Bot. Jahrb. Sysr, 83: 167 (1964).—

Pteropbylla fraxinea D. Don, Edinburgh New Philos.

J. 9: 93 (1830): Walp., Repcrt. Bot. Syst. 2: 374

(1843), 5: 834 (1846); Midi. Berol., in Walp.. Ann.

Bot. Syst. 5: 31 (1858). — Windmannia fraxinea (D.

Don) Kuntze, Revis. Gen. PI. 1; 228 (1891).— Type;

C. Smith s. n. (Moluccasi, Honimoa, Apr. 1797 (holo-

, LINN-SM!).

Artioldia heterophylla Blunte, Bijdr. Fl. Ned. Ind.: 869

(1826); DC, Prodr. 4: 12 (1830); G. Don, Gen.

Hist. 3: 20J (1834); D. Dietr., Syn, PI. 2: 1313

(1840); non Weinmannia heterophylla Kunth

(1823). Weinmannia sundana Miq,. Fl. Ned. Ind.

1 (1): 718 (1856); Engl.. Linnaea 36: 639 (1870),

Nat. Pflanzenfam., ed. 2, 18a: 256 (1930); SchefE,

Nat. Tijdschr. Ned. Ind. Batavia 34: 38 (1874);

Koord. 8c Valeton, Bijdr. Boomsoort. )ava; 403

(1900); Backcr, Schoolfl Java: 472 (1911); dcWii,

Rumnhius Memorial vol.: 411 (1959).—

Winatnannia sundana (Miq.) Kuntze, Revis. Gen.

PI. 1: 228 (1891).—Type (‘ in Javae sylvis montanis

prov. Batam”, probably in error): lecto-, Itère desig-

nated, chosen bv FloOGI AND, Reinwardt s.n.,

Amboina, July 1821,1.! [Hcrb. Ludg. Bat. 908.239-

...452|.

Spiraea pinnata Blume, Catalalogus: 76 (1823); Nees,

Flora 8: 148 (1825); non Moench. (1794).—

Arnoldia pinnata Blume, Bijdr. Fl. Ned. Ind. (15):

868 (1826); DC., Prodr. 4: 12 (1830); G. Don,

Gen. Hist. 3: 201 (1834); Hassk., PL Jav. Rar.: 324

(1848); Blume. Flora 41: 254 (1858); non

Weinmannia pinnata L„ (1759).— Weinmannia hlum-

ci Planch., Lorid J. Bot 6: 470 (1847); Miq., Fl.

Ned. Ind. 1(1); 718 (1856); Engl., Linnaea 36: 640

(1870). Engl., N3t. Pflanzenfam., ed. 2, 18a: 256

(1930); SchefE, Nat. Tijdschr. Ned. Ind. Batavia 34:

39 (1874); King, Mat. Fl. Malay Penins. 9: 299

(1897); Koord. & Valctun, Bijdr. Boomsoort. Java:

399 (1900); Pamp., Ann. Bot. (Rome) 2: 89 (1905);

Koord.-Schum-, Syst. Verz. 1, Fam, 120: 4 (1913);

Backer, Schoolfl Java* 472 (1911); Koord., Fxkurs.-

Fl. Java: 311 (1911), Koord. &L Valeton, Atlas

Baumatt Java I: t 200 (1913); Ridl., Fl. Malay.

Penins I: 682 (1922); Corner, Wayside Trees

Malaya I: 201, t. 52 (ed I, 1940; ed. 2, 1952), 1:

224, t. 55 (ed. 3, 1988); Backer & Bakh. f„ Fl. Java

1: 506 (1963); Bernardi, Bot. Jahrb. Sysr. 83: 161

(1964); Whitmorc. Guide For. Brit. Sol. lsl.: 59,

133, 152, 206 (1966); Stcenis. Mount. Fl. lava: t.

13, fig. 7 (1972): Whitmofe Tree Fl. Malaya 1 179

(1972 ).—Weinmannia arnoldia A. Gray, LES. Expi.

Exped., Phan. 1: 675 (1854 ).—Windmannia blurnei

(Planch ) Kuntze, Revis. Gen. PI. I 228 (1891).—

Type (Crescit in sylvis montanis Javae insulae):

lecto-, here designated, chosen by HOOCI AND,

Blume 816a, 12 [Hcrb. Ludg. Bat. 908.238-,..539];

syn. non.

Weinmannia hursfieldii Miq., Fl. Ned. Ind. 1(1): 718

(1856); Engl., Linnaea 36; 642 (1870); SchefE, Nat.

Tijdschr, Ned. Ind. Batavia: 39 (1874).—

Windmannia hvrsjieldii (Miq.) Kuntze, Revis. Gen.

PI, 1: 228 (1891).—Type (Java, bij Soerabaja, door

Horsfleld ontdekr): Harsfteld s.n , (holo-, BM!); syn.

nov.

Arnoldia fraxinifolia Blume, Flora 41: 254 (1858);

ADANSONIA. ser, 3 • 1998 • 20(1)

Hopkins H.C.F.

Miill. Berol., in Walp., Ann. Bot. Syst. 5: 31

(1858).— Weinmannia fraximfitLia (Blume) Miq., Fl.

Ned. Intl. 1(1): 1095 (1858): Engl., Nat.

Pflanzenfam., cd. 2, 18a: 255 (1930).—Type (ln

sylvis elatioribus Javae occidentalis): lecro-, here desig-

nated, chosen by HuOGlAND, Blume t. n. , Java, in

cacuinine monvis, Malabar, L.! [Hcrb. Ludg. Bat.

908.238-,..577 & 908.238-...578]; >yn. nov.

Weinmannia papuana Schltr., Bot. [ahrb. Syst. 52:

162 (1914); Engl., Nat. Pflanzenfam., ed, 2, 18a:

256 (1930),—Type: Ledermann 9784, North East

New Guinea, in dense hnmid forest on the April

River, 200-400 m, Nov. 1912 (holo-. B; iso-, L!);

syn. nov.

Weinmannia («dtrmannii Schltr.. Bot. Jahrb. Syst 52:

162 (1914): Lane-Poole, Rep. For. Res. Terr. Papua

New Guinea: 63, 90. 91 (1925); Engl., Nat.

Pflanzenfam., ed. 2, 18a: 256 (1930): L.M. Perry, J.

Arnold Arbor. 30: 160 (1949).—Type: Ledermann

9922 , North East New Guinea, in montane forest

on Lordberge, ta. 1000 m, Nov. 1912 (holo-, B: iso-,

K!, L!); syn. nov.

Weinmannia romentelbi Schltr,, Bot. Jahrb. Syst. 52:

163 (1914); Engl.. Nat. Pflanzenfam., ed. 2, 18a:

256 (1930); Bemctrdi, Bot. Jahrb. Syst. 83: 179,

t. 28 (1964).—Type: Ledermann 8172 (not 8173 as

given by ScHLECllTfcR [1914]), North East New

Guinea, in forest at foot of Hunstein peak, 200-

400 ni, Aug. 1912 (holo-, B!); syn. nov.

Weinmannia blumei Planch. var. major Ridl., Fl.

Malay. Penins. 5: 307 (1925).—Type: Ridley 16029 ,

Malay Peninsula, Pahang, Gunong Tahan, ca. 5500

ft., July 191 1 (lecto-, here designated, chosen by

Hoogland, SINGÎ; isolecto-, BM!, K!, SING!); syn.

nov.

Weinmannia dictyoneura Schltr., Bot. Jahrb. Syst. 52:

163 (1914), non Diels (1906).— Weinmannia alla

Engl., Nat. Pflanz.eniatn., ed. 2, 18a: 256 (1930);

Bernardi, Boc. Jahrb. Syst. 83: 160 (1964).—Type:

Ledermann 10129 , North East New Guinea, in

montane foresr on l ordberge, ca. 1000 tn, Dec.

1912 (holo-. B; iso-, K!); syn. nov.

Weinmannia borneensù Engl., Nat. Pflanzenfam., ed.

2, 18a: 256 (1930): Airy Shaw, Bull. Mise, lnt'orm.

1940: 259 (1940); Heine, Mirt. Bot. Sta.ussamml.

München 1: 212 (1953): Bernardi, Bot. Jahib. Syst.

83: 164, t. 17 (1964).— Type: Native Collectât 722,

Sarawak, s.loc., s.d. [fidc spécimen in PNII: road to

summit of Gunong Matang, 12 Aug. 1911, leg.

Dabong] (holo-, B; iso-, BM!, BO!, K!. L!, PNH);

syn. non,

Weinmannia dnlitcnsis Airy Shaw, Bull. Mise, lnform.

1940; 259 (1940).—Type: Native Collector 1671 ,

Sarawak, Dulit Ridge, forest nr. waterfall, ca.

1200 m, 10 Sep. 1932 (holo-. K!: iso-, L!, SING!);

syn. nov.

Weinmannia hypoglauca Kanch. & Hatus., Bot. Mag.

(Tokyo) 56: 111, t. 7 (1942).—Type: Kanehira &

Hatusima 12797 (Irian Jaya), Boemi, 40 km inward

of Nabire, 400 m, 11 Mar. 1940 (holo-, FU, photo

at K!; iso-, BO!, L!); syn. nov.

Cortexpapeiarius Rumph., Herb. Amboin. 3: 212,

t. 137 (1743), nom, illeg.

Small to large tree, up to 25(-40) m high.

Young stems pubemlent to romentose, rarely glab-

rous, older stems glabrescent with numerous

lenticels: branching not usually dichotomous.

Stipules pften caducous, ± orbkqlar to subreni-

form to broadly spathulate, 0.8 X 0.8 to 1.5 X

1.8 cm, narrowed at base, apex broadly rounded,

abaxial surface shordy strigose especially towards

base, adaxial surface glabrous, Leaves imparipin-

nate, with (O-)l-S pairs of latéral leaflers. total

lengtb 5.5-15.5 cm including petiole of 1-3 cm;

rachis segments 1-2.5 cm long; petiole and rachis

segments terete, indumenrum varying from glab¬

rous to totnentose-velutinous on adaxial side or

abaxial side or both; latéral leaflets ± sessile,

usually broader towards the base, varying from

lanceolate to narrowly elliptical to narrowly ovate

to ovate, the largest per leaf (2.2-)4.2-8.5(-12) x

(0.8-)1.2-3.5(-4.5) cm, proximal leaflets usually

smaller rhan more distal ones, base frequently

uneciual to dimidiatus, rounded ro cuneate on

either side, apex acuce to acuminate; terminal

leaflet narrowly elliptical to narrowly ovate, 2.5-

10 x 1-3.3 cm including petioiule of 0.3-1.5 cm,

blade t same size as largest latéral leaflets or

larger, base attenuate into petioiule, apex acumin¬

ate: blades chartaceous to subcoriaceous, glab¬

rous, not punctate below; margin crenate, 8-14

notches on each side in the latéral leaflets, crena-

tions rounded and acroscopic or triangular and

not acroscopic; midrib prominent and shortly

hirsute below, indented altove and sometimes

hirsute towards base; secondary and tertiary veins

± Hat above and minutcly prominent below,

sometimes drying paler than intervenium.

Inflorescence 1-3 opposite pairs of latéral dyads

or tetrads at most distal node of main stem,

inserted in sériés when > 1 pair; apical bud o( the

main stem between the pednncles of the central

dyads densely sericeous; bud at apex of peduucles

in angle between the racemcs sericeous; some¬

times a few successive nodes on ont" shoot prod-

ucing partial inflorescences simultaneously;

peduncles 0.3-1.8 cm long; racemes 7-5-15 cm

ADANSONIA, sér. 3 • 1998 • 20 (1)

Weinmannia in western Maiesia

Fig. 7.— Weinmannia traxinea A, flowering shoot showing an inflorescence composed of 4 partial inflorescences (mixture of dyads

and tetrads): note dormant apical bud and buds wlthin each partial inflorescence; B. latéral leaflet, fypical of W. Itaxinea " dulitensis

C, latéral leaflet, typical of W. fraxinea "bomeensis'; D, latéral leaflet, typical of * blumei"; E, latéral leaflet, typical of W. traxinea from

the Moluccas; F. bisexual flower; G. female flower, one petal removed; H. male tlower; J, debisced capsule, central column not évi¬

dent; K, seed. (A, Hopkins S Bradford 5006; B, Hopkins & Bradford 5013; C, Meijer SAN 22106; D, Nousjet 670 ; E, Kuswata &

Soepadmo 250; F, J, K, Dickison 21 5; G, Geesink 8953 ; H, Hopkins <S Bradford 5006). — Drawn by C. Monniê.

ADANSONIA. sér. 3 ■ 1998 ■ 20(1)

Hopkins H.C.F.

long; peduncles and axes of racemes minutely

puberulenr to tomcntose. Floral buds inserted in

fascicles; floral bracts ligulate-carinate 0.5-1 mm

long, caducous, wich short flattened hairs.

Flowers uniscxual or bisexual; pedicel (1 1-J1.5-

3 mm long, minutely hairy; calyx lobes triangu-

lar, 0.6-0.9 mm long, hirsute at base; petals

oblong to obovate. 1.1 -1.8 X 0.7-1,2 mm, apex

rounded or rarely ctnarginate, ciliolate or not;

dise lobes iree, oblong, 0.2-0.5 mm long, or rare¬

ly formïng an almost complété dise; m male flow-

ers: filaments ca. 2.9 mm long, anthers ca,

0.5 mm long, ovary ca. 0.5 mm, densely pubes-

cent, styles 0 1 mm, incurved; in female flowers:

filaments 0.6-1.2 mm long, anthers 0,3 mm

long, ovaiy 0.6-1.2 mm long, densely pubescent,

styles 1-1.3 mm long, straight; in bisexual flow¬

ers: filaments 3-3.5 mm long, ovary 0.6-0.8 mm

long, pubescent, styles 1 5-2 mm long.

Capsules (2-)2.5-4(-6) X 1.5-2(-3.1) mm at

dehiscencc, plus styles 1 -2 mm long, the exocarp

pubescent; calyx lobes usually persistent; central

column weakly developed. Seeds 0.8-1.1 mm

long, comose at both ends, the haits up to 2 mm

long.—Fig. 2A,C,D; 7A-K; 8.

JUVENILE FOUACE. —Small seedlings hâve 3-5

pairs of chartaceous leaflets tinged with red, the

margin dentate and the teeth acroscopic; the

stipules are persistent, ± round and ofien toothed

(Hopkins & Bradford 5017, Bornéo). Older seed¬

lings hâve up to 12 pairs of leaflets with similar

margins and the stipules are again often persis¬

tent and toothed (Hopkins & Bradford 5003,

Bornéo; Verheijen 4397, Flores, Steenis 7398,

Java p.p). Indumentum on the main stem and

leaf rachises varies from almost glabrous to

tomentose.

Table 1.—The number ot herbarium specimens with flowers of different morphological types, as determined by the

relative lengths of the styles and filaments, in Weinmannia fraxlnea from various régions and two closely related

species Sex of flowers was determined at anthesis (columns 2-4) or rarely in fruit (column 5) but never prior to élong¬

ation of the filaments.

Geographical

région

Male

flowers

Bisexual

flowers

Female

flowers

Fruit with remnants

of tiny stamens

(i.e. from female flowers)

Breeding System

(as indicated by

morphology of flowers)

Sumatra

dioeclous

Malay Peninsula

largely dioecious

Java

1 + 3 (male

+ bisexual)

polygamodioecious

Bornéo

partly dioecious

Lesser Sunda

Islands

dioecious

Moluccas

dioecious

New Guinea

largely hermaphroditic

Solomons

hermaphroditic

Sulawesi

( W. devogelii, see

part 2, p. 48)

dioecious

Vanuatu

( W. maegillivrayi,

see part 3, p. 85)

2 (male

+ bisexual)

polygamodioecious

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in western Malesia

BrhediNG SYSTEM. —Varying with geography,

either dioecious, polygamodioecious or herma-

phroditic. See Table 1.

FlELD NOTES. —Small, often shrub-like tree 5-

15 m tall at high altitude, to tall, siender tree up

to 25(-40) m high X 50 cm dbh at lower altitude.

Buttresses usualiy absent; when présent, up to

2 m, concave, rounded. Bark variable but ftc-

quently brownish and llaking; varying from dark

brown, pink- or orange-brown, greyish, fawn,

reddish purple to rich tawny rust, and the texture

scaling, fissured, smooth, llaking or rugose-ienti-

cellate. Inner bark variable, often reddish; also

orange-yellow, brownish grey, golden-brown or

reddish brown and tibfous. Slash wood dirty yell-

ow-white; no exudatc. Hearr wood light orange

to beefy rcd. Stipules sometimes prominent;

young leaves, stems and stipules often pinkish,

bright red or purplish. Mature leaves shining

above, sometimes with red margins, the rachis

brown-red, the stipules light green; old leaves

bright red. Flower buds green, yellow, pink or

red; inflorescence axes green or dull red; pedicels

sometimes red. Flowers white, yellowish, pinkish

or pale green; pedicels and calyx greenish or pink¬

ish; pelais and filaments white or creamy pink;

dise yellowish; anthers white to sulphur; ovary

red with styles pink or white, the hairs white; flo-

wers lightly scented or not. Young fruit green,

reddish green, yellow, bright red, purplish or red-

brown; old fruits blackish, seeds yellowish white.

Distribution and ec.ol.ogy. — From northern

Sumatra eastwards chrough Malesia (Malay

Peninsula, Greater and Lesser Sunda Islands,

Moluccas, New Guinea and Bismarck

Archipelago) to the Solomon Islands, absent

from Sulawesi and the Philippines. (0-)500-

2000(-2700) m. In western Malesia, usualiy from

mid to high élévations and more frequent at

lower élévations in the eastern part of its range

(Malay Peninsula: (450-) 1000-2065 m; Sumatra:

500-2700 m; Java: 600-2400 m; Bornéo:

(0-)500-1970 m: Lesser Sunda Islands: 500-

1500+ m: New Guinea: 10-1450(-2250) m;

Solomons; 10-620 m).

In western Malesia, at lower élévations in pri-

mary and secondary forest, includirtg with

Casuarina. Locally common small tree above

500 m, in hill forest, often on ridges and spurs or

on siopes below ridge top. Varying in abundance

from uncommon to one of main constituents of

montane forest (e.g. Jacobs 8201, Sumatra),

sometimes locally common and clumped. At

higher altitude found in mossy heath forest,

montane rain forest and dwarf, montane scrub

and alpine shrubbery Often on young volcanic

soils and open, stony ground, sometimes recently

burnt and dominated by ferns. Also grows on

soils derivcd from sandstone, daciic and ultra-

basics and recorded from acidic, water-logged

sands in Agathis forest ( Kostermam 12903),

sandy, kerangas-like soil, and mixed peat swamp

forest at sca level (Bornéo).

Sr.LECTED COLLECTIONS (from a total of 347 scen for

this région).—SlIMATRA: Aceh: Steenis 9543, Gayo

lands, Gn. Kemiri, 1800-2000 ni, st., 6 Mar. 1937

(BO); Wilde & WUde-Duyjjes 13087 , Gn. Leuser Nat.

Res., Gn. Baudahara, track from kamp. SeldokNL to

blang S of suffltnit, 25 km NNW ol Kutatjane,

1800 m, yfl., 20 Julie 1972 (K, KRP, L); Wilde &

Wilde-DuyJjes 14151, Gn. Leuser Nat. Res , Gn.

Ketambe, 8-15 km SW from mouth of Lan Kerambe,

40 km NW of Kutatjane, 2000 m, ll„ 9 Aug. 1972

(K, KEP, L): Wilde & Wilde-Duyfies 16638, Gn.

Leuser N 31 . Res.. Gn. Marnas, 9 km SW from mouth

of Lau Ketambe. 30 km NW of Kutatjane, 1500 m,

fl,. 6 May 1975 (K, KEP. L). {Tapanuli 1; Alston

14891, Baniara, S of Sidikal.ing, fl., h., 28 Mar. 1954

(BM, L); Marernitn 6, bb 5261, Toba plateau, nr.

Pansurbatu, 2693 m, fl., 24 |an. 1923 (BO, E). W

Sumatra: Abid gl Malin Maradjo 6, bb 18608, nr.

Kcrintiji Tengah, 1400 m, st-, 20 Mar. 1934 (BO);

Bürmemeyer 908, NW dope Talaman, Ophîr Distr.,

1900 m. fl , 28 May 1917 (BO); Meijo 4218, Mt.

Singgalang nr. Bulcittinggi, 2000-2200 m, fl., yfr., 1

July 1956 (L); Meijer 5898, Mt. Sago nr.

Payakumbuh, 1500 m, yfr., 22 J une 1957 ( L). (E

coast): Bangham dr Bnngham 1020, I r.nl from Mcdan

to top of Sibajak volcano, 4200 6500 fi., fl., 15 Feb.

1932 (K, SING); Liirsing 7941, Central Habinsatan,

righr bank of Kuwal between Parsoburan and Nassau,

850 m, buds, 15 Nov. 1920 (BO, L); leysmun s.n.,

Lingga, Mt. Tanda, st. (BO, L); Yates 2007, Braxragi,

4500 fr., fl., yfr.. 12 Mar 1926 (BO, L, SING).

lienkulu : Mesaerip 20, bb 8753, Kroë, nr. Gn.

Kewala, 800 m, st., 7 Mar. 1925 (BO); Vottgd bb

17032, Lebong, nr. Bt. Daun, 1800 m, st., 1 Apr.

1932 (BO). Palembang. Steenis 3617, Gn. Pesagi. N

ridge SSE ot L. Ranau, 2000 m, st., 4 Nov. 1929

(BO). Lampong: fucobs 8201. Mt. Tanggamus,

5 e i6’5-t04 ft 41'E, 1500 m, fl., 1 May 1968 (A, BiSH,

K, KEP, L).—Peninsular Malaysia: Kedah. Wyatt-

Srnitb KEP 79272, top of Kedah Peak, 3950 ft., fi., 28

Apr. 1957 (K, KEP, L). Perak-, Ridley 3027, top of

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

Gn. Hijau, fl., Mar. 1892 (BM, K, SING); Wrav

3814, Gn. Bahu, Larut, 5000 fi., fr., Mar. 1890 (K,

P, SING). Kelantan: Symingtan KEP 37700, Gn.

Stang, st., 10 Oct. 1934 (SING). Trengganu:

Cockburn FRI I0S28, E approach ridge Gn. Mandi

Angin, 4200 ft., budv. 12 ]uly 1962 (K, KEP, L):

Moysey dr KiahSE31040, Gn. Padang. 4000 fc., buds,

June 1937 (KEPv SJNG). Selangor. Maxwell 78-318,

Gendng Highlands, Gn Ulu Kalt, 1700 m, fl., 3 June

1978 (El. Pahang. Corner SF 33161 , Frascr’s Hiil,

4000 ft., fl., 10 Aug. 1937 (A, BO, E, SING); Klnss

126 or SE 12124, Gn. l'alun, Padang Sebrane, 4900

ft., fr., 6 July 1923 (SING); Soepadnio 050, Éetween

Wray’s camp and thc Padang, l’aman Negara, 3000-

5000 ft,, fl., 4 Sep. 1970 (A, K, E, SAN, SAR);

Syming/ur/ S F? 36224, Cameron Highlands, summit

of Gn. Tcrbakar, fr., 12 Api. 1934 (K. KEP, SING);

Wbiimore FRI 3366, main NE ridge of Gn. Rebom,

Krau Game Reserve boundary, 5000 ft., fr., 18 Mar.

1967 (A, K, KEP, L) Malacca: Griffith 2507, Gn.

Toondot ledaiig top, fl,, 1861-62 (K, P); Ridlty

3295, Mr. Onhir, buds, June 1892 (K, SING).

Johore Mat Àsri bin Ngah Sanah FRI 38681, Gn.

Ledang, nr. lelecom building, 450 m, fl., 20 Apr.

1993 (KEP): Whitmore FRI 12394. NW Johore, Gn.

Ledang (Mt. Ophir), 3700 ft., fl., 17 July 1969 (A, K,

KEP, L).—Java (W = vvesr, C - central, E = cast):

Arens 79, E, Malang, Ardjuna, between Kokopan and

Lalijiwo, 2400 m, fl,, 15 Occ. 1915 (BO. L); Backer

7118, W, Bantarn, Gn. Aseupan nr. Menes, 400-

600 m. fr., 12 Mar. 1913 (BO); Backer 22969, W,

Bogor, Gn. Beser, nr. Tidadap, S of Tibeber, 1050 in.

buds, 10 Sep. 1917 (BO); Backer37408, E, Malang,

Pasuruan, Gn. Ardjuna, above [ jeter, 2000 ni, fl., 3

Oct. 1927 (L); Balgooy & Wlriadmata 2913, W,

Nirmala Estate, Gn. Halimun area, 1350 m, fl., 10

June 1980 (A, L); Boschpmefitation 6691, E, Madiun,

Kedunggalar forest, fr., Mar. 1927 (BO); Burck 483 ,

W, Priangan, Garut/Karotjok, buds, ft., 4 July 1891

(BO, L); Hnçhreutiner 1941, W, Bogor, Mt. Salak,

towards W on Gn. Bunder, 800 m, fl., 2 Oct. 1904

(L); Junghuhn s.n.. G, Semarang, Medinie, Ungarang,

yfr. (L); Kalshoven VI, W, Bogor, Tjiadhu (Cin.

Salak), 900 m. 11., Nov. 1918 (BO); Kaordcrs 7640,

W, Bogor, Tjibodas, Tjiputri, fl., 22 Fcb. 1890 (BO,

L); Koorders 7645, W. Bogor. Takoka, 1100 m. fl., 3

Aug. 1890 (BO, L, P); Koorders 6746, W, Priangan,

NW Galunggung, nr. top Pangcnijongan-

Telagabodas, 2229 m, 11., 1 Jan. 1891 (L); Koorders

7648 , W, Bantam, Gn. Pulasari, Kilmdjan,

Pandeglang, 1050 m, lt., 13 June 1892 (BO, l );

Koorders 7652, C, Semarang.Telomjo, Ambarawa, fl.,

fr., 9 Oct. 1888 (BO, K, L); Koorders 7653, C,

Banyumas, nr. Pringotnbo, Bandjarnegara, yfr., 9

Nov. 1891 (BO); Koorders 7657, C, Banyumas,

Midangan Mrs. nr. Pringombo, 700-1000 m. fl., 15

Nov. 1891 (BO, L); Koorders 12441, E, Madiun, W

Gn. Wilis, Ngebel, 1350 m, yfr., 22 Oct. 1892 (A,

BO, L, SING); Koorders 26546, W, Priangan, Garut,

Wanaradja, Pasir, Pasanggrahan Pangontjongan, fr., 5

Eeb. 1897 (BO, L); Koorders 27952, C, Semarang,

Ambarawa, Telomojo, fl., 13 June 1897 (BO, L);

Koorders 34183, E, Madiun, Ngebel, Ponorogo, fr.,

19 Nov. 1900 (BO, L); Koorders 37915, E, Malang,

I engger, 1650 ni, fl., 30 Oct. 1899 (BO, K); Koorders

38229, E, Malang, Gn. Ard|Uno, Pasuruan, 2000 m,

fr.. Il Nov. 1899 (BO, K L); Lorzing 716, C, Tlerep,

Prahu, N Sendara, 2000 m, buds, 24 Nov. 1912 (BO,

I ); Steenis 5110, W, Bogor, TjiapuskJoof, Gn. Salak,

800-1000 m, IL, 31 July 1932 (A, BO, K, L, SING);

Steenis 17615, W, Bogor. Gn. Pangrango. 2500 m,

st„ 29 May 1950 (BO); Winckcl 1559, W. Bogor,

I jadasmalang, nr. Tjidadap, S of Tjibeber, 1000 m,

fl. fr.. 11 Aug. 1923 (BO. K E, SING).— Bornéo:

Saiiaii: Ahan Gibot SAN 60723, Distr. Ranau. Kamp.

Kilimu Ranau, 2.5 miles E of Ranau along Poring Rd,

2000 ft„ fr., 18 Sep. 1967 (BISH, SAN); Aban Gibot

SAN 62024, Distr. Tuaran, Kampong Parad, fr., 18

May 1968 (L, SAN, SAR); Aban Gibot SAN 66831,

Distr. Ranau, Copper Mining Area, Mamut, buds, 2

May 1970 (L. SAN); Aban Gibot SAN 95210, Distr.

Beluran, Ri. Monkobo, 1950 m, st„ 15 Mar. 1982

(SAN); Alabazo SH (SAN) A 3626, for est Distr.

Jambunah, Kg Zanateir. 2500 ft., fr. 26 Mar. 1952

(L. SING); Carr SF27228, Mr. Kinabalu, nr. Bundu

Tuhan, 4000 ft.. fl., .3 May 1933 (SING); Chew &

Corner RSBN 4398, Ml. Kinabalu, Bembangan R.,

5400 ft., st., 20 Eeb. 1964 (K, L, SAN); tSiemens &

Cletncns 50693, Mc. Kinabalu, Gnrulau Stnir, 5500

ft., fr., 4 Dec. 1933 (A, BM); Cletncns cr Clemens

51173, Mt. Kinabalu, Penibukan ridge, 4000 ft.. fl., 6

Nov. 1933 (A, BM. K); Cockburn SAN 84928 , Distr.

Eahad Ditu. Summit trail of Gn. Tribulation, Sg.

Segama, 900 m, fr., 18 Aug. 1976 (K. !.. SAN. SAR);

Rfadani SAN 89394, Distr. Ranau, fom Bt. Ampuan,

2500 ft., fl., 18 Nov. 1978 (L, SAN, SAR); Meijer

SAN 51588, Distr- Sandakan, Mt. Melian nr. Kiabu

Labuk, 2800 ft., juv., 14 May 1965 (SAN); Mikil

SAN 31867, Distr Tantbunan, I rusmadi above

Kionop, juv., 10 Oct. 1962 (KEP); Mikil SAN37765,

Distr. Penumpang, .Suran Trail, 890 ft., fr., 15 Dec.

1963 (L, SAN, SAR); Sorte SAN28510, Distr. Ranau,

spur of Kg Tamis Tenompaok b.R., 4700 ft.. yfr., 19

Fcb. 1962 (K, KEP. E, SAN, SAR); Sundaling SAN

129697. Distr. Lahad Datu. FR Malau, fl., 30 July

1990 (SAN).— Brunei: Ashron BRUN 1044, Bt.

Pagon ridge, 4750 ft., fr., Mar. 1958 (K, L, SAR). —

SaRawaK: Ashtott S 21116, Bt. Lumut, Carapa Amau,

Glu Mujongv Hose Mts., 950 m st,, 14 Apr. 1964

(K, SAR); Chai S 18535, Gn. Matang, 800 m, fl., 31

July 1963 (A, K, KEP, L. SAN. SAR); Chai S35503,

Perupayang nr. foot of Apo Duai Range, Kelabit

Highlands, Baram Distr,, 4tli Div., 1180 m, yfr., 14

Nov. 1974 (A. K. E. SAN); Dyg. Awa & l.ee S 51142,

Tama Abu Range, Bario, 4th Div., 1750 m, fl., 19

May 1988 (K, KEP, L, SAR); Uias Paie S 26012, Ulu

Ugong, Lawas, 2800 ft., fr., 19 Sep. 1967 (K, L, SAN,

SAR); Lee S 52496, Bt. Tebunan, Ulu Tntsan, Lawas,

ADANSONIA. sér 3 • 1998 • 20(1)

Hopkins H.C.F.

5th Div., 1450 m, fl., 9 May 1986 (K. L, SAN, SAR);

Martin S 37053, Gn. Mtilu National Park, Tutoh,

Baram, 4th Div., 1310 m, 11., 24 Fcb. 1976 (K, L,

SAN, SAR); Murthy & Ashton S 22621, Usun Apau,

Long Nyaiau, Dapoi, Tinjar, 700 m, st., 4 Apr. 1965

(A, K, L, SAN, SAR); Nuoteboum & Chai 1696,

Kalabit Highlands, Bario, daka sina kaja hill, 1000 m,

yfr., 25 Mar. 1970 (BISH, K, Kbl\ L. SAR); Pickles S

3777, Mt. Kalulong, 3"l4’N-I l4°4l'E, 4310 ft. fr.,

19 Feb, 1956 (BM. SAR).—KALIMANTAN: Enderr

3136 , W Kurei, !.. fVtak, 600 m, fl., 10 Sep. 1925

(BO, L): Geesink 8953, Kalimantan Timur, betwcen

Long Bawan and Panado, 3"52'N-115 C 42"E, 1000 m,

fl., 7July 1981 (L); Joug bb 9670, West Bornéo, Ond.

afd. Bengkajang, top Gn. Bawung, 1400 m, juv., st„

25 Mar, 1926 (BO); Kosterrnany 7540, Peak of

Balikpapan, Gn. Beratus, 1200 m, st,, 15 July 1952

(BO, K, L); Kostennans 12903, W. Kutei, Mt.

Palimasan nr. Tabang on R. BeUjan, “00 m, Fr,, 11

Sep. 1956 (R.M, K, K H P, L); Valkenbufg & Stockdak

1082, Kayan Mentarang Reserve, 2’5l'N-11 5°55'E,

950 m, fl„ 26 Nov. 1991 (A); Yaeger 203, Tanjung

Puting, Kalimantan 1 engab, 0 m, st., 16 Nov. 1985

(A).—LESSER Sunda IstANDS: Bail: Kos ferma ns et al.

KK&SS 162 , Bedugul forest région, Mt. Batukan

complex, 1300 m, IL, 25 June 1958 (A, BM, BO, K,

L, P); Meijer 10400, Mc. Aguns, 8°20'S-115°30'E,

st., 17 June 1976 (L); Seujf 17, bb 17276, Klucnkung,

nr. Pengedjaran, 1200 m, fr., 22 Feb. 1933 (A, BO);

Steenis 8043, Gn. Ahang, 1500-2000 m, st., 9 Apr.

1936, (BO). Sumbatva: Kostermans 18439, W

Sumbawa, Mt. Battilanrch, Brangbossang, trail front

Batudulartg to Pusu. 800-900 m, st., 24 Apr. 1961

(K, L). Sumba: Verbetjen 4028. s.loc., buds (L);

Voogd 1901, W Sumba, 500 m, fl., 30 Nov. 1934

(BO, L). Flores-, Hooghtnd 12646, W Flores, Poco

Raneka nr. Ruteng, 1400 m, fl., 25 June 1975

(CANB, L): Kostermans 22091, W Flores, SE part

Mbengen, 600 ni, buds, 10 May 1965 (K, L);

Schmuta 4588, Nunang, 1100 m, fr., 2 Feb. 1980 (L);

Verheijen 4360, Karat-Lelak, st., 16 July 19"4 (L).—

MOLUCCA5: Morotai Kostermans 1097, Gn. Parc 2,

900 m, 26 May 1949, (BO, L). Bacon Haan 2, bb

23122, nr. Masurung. 500 m. sr., 12 Aug. 1937 (A,

BO, L). Sidti Jsls.: Atjèh (Fxp. v. Hulstijn) 383, Eil.

Sulabesi (~ Sanana), Gn. Patahoi, IL, 1913-14 (BO,

L). Seront: Burley et al. 4363, 40 knt E of Masohi in

Wae Rutan/Wae Ruwata (Ruwa) catchment areas,

150 m, fl„ 7 Dec. 1990 (A, K. KEP, L, P);. Runen

1649, W Ceram, W of Piru, 0-100 tn, fl., 25 Sep.

1918 (BO, K, L); Runen 2221 , C Serarn, H Elate S of

Railiu, 400 m, 11 , 13 May 1919 (BO, L). Ambon:

Burlage 253, Gn. Tuna, fl., 17 July 1900 (BO);

Dumont d'Urville 1311, s.Loc., fl., Ôct. 1823 (P);

Kornassi 1181, Gn. Harumesen, 0-100 m. fl., 25 Apr.

1918 (BO, K, L); KusivaUt d Soepadrno 250, Wa-ai,

slope of Mc. Salhutu. 100-200 m, fr., 7 July 1959 (A,

BISH, BM, K, KEP, L, P); Matatula 5, bb '14261, nr.

Puta, kamp. Hatu, 250 m, st„ 14 Dec. 1929 (BO);

Rant 112, Karang panjang, behind Soja rd., fr„ 30

May 1929 (BO); Vriese cr Teysmann s.n., s.loc., fl.,

1859-1860 (L); Wiljes-Hissink 21, foorhills of Salahutu

Mt. nr. Wae, 200 m, buds, Apr. 1948 (A); Zippel

72d, s.loc., fr., Sep. 1828 (L). Uli Asser Jsls.: Malessij

4, bb 14303. Haruku, nr. Taënenitu, 300 m, fl., 20

Mar. 1930 (BO). Banda: s.colf, s.n., s.loc.. fl. (L).

Cuit, in Hort. Bog, ex Moluceosr. Fortnan 51, J130,

fr., 25 Feb. 1956 (A, BO, K, L); Jacobs s.tt, J130a,

male fl., 12 Mar. 1958 (BM. K, L, P),

Local names. — Sumatra: Antahasi or Atahasi

(Batik language, ’foba), Bocnoha (Toba), Resak kero

(Malay language), Bocnga I-octih (Lampung lang.),

Bringin randock (language?).—MAL. PEN.: Kasai

bukic (Burbill & Holttum SF 6853), Scnrang

( Wbitmore FRI 3881), Pa'ang (Bidin CF 4200 ).—

JAVA: Sundanese: Ki merik(h), Karo tjok, Ki papa-

tong, Kihiris, Kiringït, Tjirome, J jeremeh, Ki

Tjercrne, Ki nicong. Angrict pissang, Kibiroe, Ki

adjae, Ki-papatrong. Javanense: Dali kling or

Dalilceling, Wlingi, Kelis, Panejalkidang, Sampar

kc-dang, Gagar, Kerjemé, Serak, Lere, Gringging,

Damaran. ?Klis, :djeng gïtri, Nagasari gunung,

Sap(i)enan, Sadan, Tembatang or Têinbagan,

S(e)riwaJik, Wuru kunjet. Language not specified:

Damaran [Bat/ter 16223), Ki soeren goenoeng

( Konrdcrs 26.546), Legarie Tjantigi (Kalshaven VI),

Pasang kengkeng [Voogd 721), Kajoe djaran [Winckel

1963). —BornfO: Tekaran(u) (Kelabit). Uban (Tban),

Tansang t.ang (Iban), tkaran [Nootebnom & Chai

1696), Kayu-papan (Murut).— Lesser Sunda

lSLANDS: Kémêrakdtlan (Balineese lang.). Larang

(Flores).—MoLUCCAS: Mjrah. k - Doka2ra = Sela, k

(Tobela lang., Bacan), Daon mejang (Ambon),

Kahoenar (Malay lang., Ambon), Lagoendi hoetan

(Malay, Ambon dialect), Taheroe (Malay. Ambon

dialeet), Meongmerah (Malay, Ambon dialect),

Mooeroca (Haruku lang.).

LOCAI. USES. — BORNEO: The leaves are used for

dying cloth; rhey are boiled in water and then

mixed wich clay to makc a blackish dye ( Chai S

35503). Firewood.

Nomenclature and typification.—

HOOGEAND worked extensively on the nomen¬

clature and typification of Weinrnannia Jraxinea

and its synonyms. Several lectotypes thar he had

chosen are formally designated here.

Under article 42.1 of the International Code of

Botanical Nomenclature (GREUTER 1994),

Pterophyllu Jraxinea D. Don, rhe basionym for

Weinrnannia Jraxinea, was validly published in

1830 [descriptio generico-specifica) and should not

be considered a nomen nudum.

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in western Malesia

For the names published by SCHLECHTER and

placed in synonomy here, as well as for W. born-

eensis Engl., and for W ptdlei SchJtr. and W. vir-

gulata Schltr. (see HOPKINS 1998b), ihe

holotypes at Berlin hâve noi been destroycd (see

Bernardi 1964).

Taxonomy and variation. —As circumscribed

here, Weinmannia fraxinea is a widespread spe-

cies that shows a complex pattern of variation in

the number, size, shape and texture of the leaflets

and in the indumentum. Weinmannia blumei,

the naine usually applied ro rhis species in west¬

ern Malesia, is placed in synonomy wirh W.

fraxinea here (or the First rime, as are several

names from Bornéo and New Guinea that were

regarded by BERNARD! ( I 964) as distinct. The

variation in W fraxinea cannot be consistently

divided into discrète urvits either morphologically

or geographically, i.c. ir shows “checkerboard”

variation, and can thus bc regarded as an ochlo-

species (CRONK, in press; WillTE 1962). Since

the variants grade into one anorher, division into

infraspedfic taxa is unwarranted. Flowever, it is

sometimes useftil to be ablc to liante variants,

especially at a local level, and in this case I sugg-

est following the example of HUXLEY & JEBH

(1993) who propose rhe use of informai nick-

names for the most distinctive variants of ochlo-

species (e.g. “ dulitensis” for the large, rubbery

leafleted form; “ blumei ” for specimens with

medium-stzed, chartaceous Icavcs, etc.). How-

ever, there are too matty intermédiare collections

for nicknamcs to bc applied consistently to ail

the materiaJ in W. fraxinea.

The majority of collections Itom throughout

the range hâve medium-sized, chartaceous to

subcoriaceous leaflets, medium-sized and usually

caducous stipules, and the indumentum on the

axes varies from puberulcnt to tomentose (Fig.

7A). The flower buds are usually pinkish and the

flowers whitish. In Sumatra, the Malay

Peninsula, Java and New Guinea, some collect¬

ions hâve sntaller leaflets (Fig. 7D) (e.g. Koorders

7646, Java; Steenis 3617 , Rappard S27 , Sumatra;

Cockbum FRI 11030, Malay Peninsula; Stevens

LAE 58123, Woods 2718, New Guinea), the larg-

est latéral leaflet per leaf excepttonally as small as

2.2 X 0.9 cm. These small-leafleted variants

usually occur at high altirude though specimens

with medium-sized leaflets sometimes occur at

similar élévation. The inflorescences tend to be

short and less well developed than in matenal

from lower élévation (i.e. fewer partial inflores¬

cences; dyads not tetrads) and the flowers are

usually whire.

Other collections from these four areas and

from Bornéo hâve exceptionally large leaflets, up

to 12 X 3.5 cm. The leaflets vary in texture from

chartaceous to coriaceous, but when coriaceous,

they often hâve minutely proiniacnr or contrast-

ingly coloured venation and a dense réticulum of

tertiary and quaternary veins. In these specimeas,

the axes are glabrous or subglabrous and the stip¬

ules are larger and more persistent. In the Malay

Peninsula niaterial like this has been named W.

blumei var. major (e.g. Ridley 16029, Wray &

Robinson 5319); in Bornéo, W. dulitensis (e.g.

Clemens 51173, Hopkins & Bradford 5013); and

in New Guinea, W. alta (e.g. Smith NGF 1355,

Lam 1574). In Sumatra, collections include de

Wilde & de Wilde-Duyffes 14151 from Gn.

Leuser, Aceh Province, flowever, the shape of the

latéral leaflets is not always the sarne, and varies

from lanceolate or ellipticai and t cuneate at rhe

base, to ovate with the base rounded and condu-

plicate (Fig. 7B). In vivo, young stems and leaves

are rubbery and difficuli to press (HOPKINS &

BradfoREi pers. obs.), In W. fraxinea u dulitensis ”

from Bornéo, the inflorescence axes are reddish,

the racemes long and robust, the flower buds

pink or bright red and the flowers pinkish at

anthesis (Fig. 2C,D).

Some other collections from Bornéo hâve ovate

latéral leaflets that are medium-sized, subcoria¬

ceous, and the base is rounded on one or both

sides but not conduplicate (Fig. 7C). They some¬

times dry black and can be densely hirsute along

the abaxial surface of the ntidrib, They corres¬

pond to W borneensis (e.g, Ampon et al SAN

71837, Aban Gibot SAN 56354, J & M.S.

Clemens 32641).

In Flores and Seram, the leaflet margins are

rather distinctly crenate (e.g. Hoogland 12646,

Schmutz 30621). Some specimens from the

islaads of Milne Bay Province, Papua New

Guinea hâve leaflets with strongly acroscopic cren-

ations (e.g. Brass 27428 and Gideon LAE 57278,

ADANSONIA, sér 3 ■ 1998 • 20(1)

Hopkins H.C.F.

both from Misima lsland) but the leaflets here

are larger than in the Lesser Sunda Islands.

In the Moluccas, rhe leaflets are typically ellipt-

ical or lanceolate and the base of latéral leaflets,

especialty the most distal pair, is strongly un-

equal, being attenuate on one skie and concave

on the other (Fig. 7E) (e.g. Ko ma su IJ SJ,

Kuswata & Soepadmo 250, Malessij bb 14281,

Teysmann s.ri., Ambon; Rutten 2221 , Seram). A

few collections from the Cycloop Mts., Irian Jaya

are rather similar (e.g. Koster BW 4302). In both

the Moluccas and New Guinea some collections

are glaucous on the underside of the leaflets (e.g.

type of W. hypoglauca from Irian Jaya; de Wiljes-

Hissink21, Ambon).

Weinmannia fraxinea is less variable than some

other ochlospecies, such as Myrmecodict tuberosa

Jack in which 16 informai entities were recognis-

ed (Huxi.l-.Y & JhBR 1993), but nevertheless it

shows many of the traits listed by CRONK (in

press) as characteristic of this type of complex

species. It has strongly polymorphie, non-hierar-

chical variation where the characters vary inde-

pendently of one another and largely

independenrly of geography and ecology. It has a

widespread distribution and shows considérable

ecological plasticiry, occurring for instance in

New Guinea from near sea level ta over 2000 m

on a variety of soil types. While distinct variants

can sometimes be recognised at a local level, they

intergrade when the whole range of the species is

considered. Similar morphologies are often

found in widely separated Iocalities and some¬

times appear to be polytopic, while other variants

occur in only one région. In some cases, at a part-

icular localiry, there can be rwo distinct variants

présent which do not intergrade, although in

another locality they do. In commun with many

ochlospecies, VP fraxinea belongs to a large genus

and has closely related, monomorphic satellites,

such as W. dcvogelii H.C Hopkins in Sulawesi

and W macgtllivmyi Seem. in Vanuatu.

Although variation is not due to interspecific

hybridization, nor does Weinmannia fraxinea

appear to be a sériés of apomictic microspecies

(see Cronk, in press), the morphology of flowers

on herbarium specimens suggests a complex and

variable breeding System (Table 1). Clearly field

observations are needed but current evidence

indicates rhat W fraxinea is predominantly dioe-

cious or polygamodioecious in some régions and

hermaphroditic in others. Successive cycles of

flowers ol different sexes on one plant, as seen in

certain Sapindaceae (VAM WELZEN 1990) and

which may occur in some of the Pacific species of

Weinmannia, would not produce the pattern that

has been observed in this species ro date. For inst¬

ance, herbarium specimens from plants in culti¬

vation at rhe Botanical Garden at Bogor, which

originated in rhe Moluccas where W. fraxinea is

morphologically dioecious, are unisexual and

provide no evidence that the trees change sex. It

seerns likely that variation in the breeding System

contributes to taxonomie complexity and it is

possible that some populations are apomictic.

The fact that distinct variants sometimes do not

intergrade where they occur together suggests a

complex breeding System and at least partial

barriers to gene flow.

CRONK (in press) discusses two hypothèses for

the origin of ochlospecies. The France hypothesis

proposes that this type of polymorphie variation

could hâve arisen in allopatrîc populations separ¬

ated front one another in réfugia during dry

periods of the Pleistocene. The geographical

région where W. fraxinea has its présent distribut¬

ion was certainly subject to considérable fluc¬

tuations in climate and sea level during this

period. In a second hypothesis, CRONK (in press)

proposes that the rapid expansion of a colonising

species with wide ecological tolérance could give

rise to the geographically unstructured variation

seen in ochlospecies. With time and outerossing,

this variation would become ecogeographically

sorred and one or ntore less-variabic species

could emerge. This hypothesis seems parricularly

apt for W. fraxinea which has wide ecological

tolérance and is often a pioneer. As suggested by

CltONK, molecular techniques could be used to

inve.stigate the genetic structure and origin of

ochlospecies.

2. Weinmannia clemensiae Steenis

J. Bot. 72: 3 (1934); Bernardi, Bot. Jahrb. Syst. 83:

166, t. 18 (1964); Cockburn, in Kinabalu Summit of

Bornéo, chapter 7: 185, fig. only (1978).—Type: J. &

ADANSONIA. sér. 3 • 1998 • 20(1)

CrjowMié

Fig. 9. —A-G Weinmannia aphanoneura A. Ilowering shoot with impaopinnale leaves. counded stipule and inflorescence of two

dyads; B, stipule; C. section ot rachis ol raceme showing ilowers inserted in fasclcles, the tloral bracts aJready failen; O, flowerbud;

E, female flower with one petal removed; F, dehisced capsule; G, seed. (A. Chew Wnn-Lek 380; B. E, Clemens 33076; C. 0, Paie

S26531: F. G. Enden 4125} — H-P Weinmannia clemensiae H, flowering stroot with imparipinnate leaves and an inflorescence of

two dyads, the apical bud continulng lo grow and productng a pair of young leaves; J, abaxial surface of a latéral leaflet: K, flower,

probably male. L, detail of K with penanth removed to show gynoeclum, dise lobes and a single stamen, M, female flower, note short

filaments; N, dehisced capsule; P, seed. (H-L, Beaman 9132\ M, Chew & Corner RSNB 4508\ N, P, Clemens 50877).— Drawn by C.

Monnié.

ADANSONIA, sér. 3 • 1998 • 20 (11

Hopkins H.C.F.

Ai.S. Clemens 27880 (Sabah), Mt. Kinabalu, in Iow

jungle near Kamborangah, 2400 ni, 7 Jan, 1932

(holo-, BO!; iso-, K! [photo at KEP], L!).

Treelet or tree, 1.5-10 m high. Young stems and

leaf rachises densely tomentose-velutinous, older

stems tomento.se becoming ± glabrous with

numerous lenticels; stems terete, relative!)' thick,

3-5 mm diameter, nodes somevvhat thickened

and leaf scars prominent. I n te modes often short,

0.5-1.7 cm. Branching noc dichotomous,

Stipules usually caducous, + orbicular, up to 1 X

1.2 cm, base narrowing, apex broadly rounded,

abaxial surface densely sericeous espectally

towards the base, adaxial surface shortly velutin-

ous. Leaves imparipinnate, with 6-13 pairs of

latéral leaflets, total length up to 13 cm includ-

ing pétiole 0.5-1 cm long; rachis segments 0,4-

0.9 cm long; petiole and rachis segments terete,

densely tomentose-velutinous, diverging front

the stem at an angle of almost 90° especially at

growing tips; leaflets bullate and carinate, the

margins strongly recurved and often rolled; latér¬

al leaflets ohlong or somevvhat ovate, the largest

per leaf 1.4-2.7 X 0.6-1 cm, leaflets towards the

base smaller, inserted ± at 90° to leaf rachis, base

equal, rounded to cordate, apex broadly acute;

terminal leaflet elliptical, 1.9-3.3 X 0.7-1 cm

including petiolule of 0.3-0.5 cm, blade scarcely

larger than largest latéral leaflets, apex acute;

blades coriaceous, the upper surface puberulent

or glabrous when the eu fiel e thick. sparsely to

densely pubescent below, the midrib sericeous;

sometitnes punctate (bases of hairs); margitt

crenate but crenations usually obscured, 5-7

notches on each side in the latéral leaflets; on

upper surface midrib indemed, secondary veins ±

at 90° to main vein, tertiary venation obscure;

midrib prominent below.

Inflorescence a pair of opposite, latéral dyads,

the apical bud of the main stem between the

peduncles of the dyads densely sericeous, often

continuing to grow vegetatively duxing flower-

ing; bud at apex of peduncle in angle between

the racemes sericeous; sometimes a few successive

nodes on une shoot producing dyads simultan-

eously; peduncles 0.3-0.7 cm long; racemes up to

10.5 cm long; peduncles and axes of racemes

densely tomentose-velutinous; inflorescences and

infructescences dense. Floral buds inserted in fasc-

icles, floral bracts obovate-carinate, with short

flattened hairs, up to 1-1.5 mm long, caducous.

Flowers unisexual (or sometimes bisexual?); pedi-

cel 1-1.5 mm long, with short erect hairs; calyx

lobes 0.8-0.9 x 0.6-0.8 mm, hirsute; pctals ob-

ovate or almost circular, 1.1-1.5 x 1-1.1 mm,

rounded at apex, margin ciliolare; dise lobes 0.4-

0,5 mm long, oblong or broadly oblong; in male

flowers' filamenrs ca. 2.5 mm long, ovary ca.

0.6 mm, densely pubescent, styles 0,1 mm,

incurved (some flowers in same inflorescence

hâve ovary 0.9 mm long and styles 0.8 mm long,

straight, and may be bisexual); in female flowers:

filaments up to 1.9 mm long, ovary 1.5-2 mm

long, densely pubescent, styles ca. 1 mm long,

straight, pubescent at base.

Capsules up to 3.7-4.5 X 1.8-2.2 mm at dehis-

cence, plus styles up to 1.5 mm long, the exocarp

densely pubescent; calyx lobes persistent; central

column weakly developed. Seeds immature, ca.

0,9 mm long, comose at both ends.—Fig. 2B,

9H-P, 10.

JUVENILE FOLIACE. —Seedling leaves: blade thin,

chartaceous, margin strongly toothed and fiat, not

recurved; latéral leaflets few (ca. 4 pairs), mid

grccu or reddish; stipules sometimes roolhed

(.Hopkins & Bradford 5015). Saplings and coppice

shoots: blade coriaceous, latéral leaflets up to 18

pairs, of similar size to chose of adult foliage but

scarcely recurved; venation indented above; mar¬

gin distinctly toothed (Hopkins & Bradford 5016).

Bkffdinc SYSTEM.— Dioecious or polvgamo-

dioccious? Few collections have flowers al an-

thesis. Spccimens with immature fruits have

short filaments, suggesting thé) bave developed

front iunctionally female flowers.

FiF.I O CHARACIERS. —Adult foliage darkish

green above, paler below with main vein brown

or pink; leaf rachises and stems brown; stipules

satiny' mid green and paler than foliage. Buds

green, white or brown front indunienium; new

growdt densely velutinous and sometimes redd¬

ish. Flowers pinkish brown or deep crealn-pink;

calyx blood red; corolla pink or pink-wliite; sta-

mens cream; whole inflorescence pink and

downy. Immature fruits pink with red styles,

infructescence axis tinged pink.

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in western Malesia

Distribu i ion and ecology.—E ndémie to

Mt. Kinabalu and Ml. Tambuyukon in Sabah.

Restricted to slunted forest on slrongly ultrama-

fic soil at ( 1640-) 1900-2600 ni. Habitats describ-

ed as low mossy and xerophylloüs scrub forest,

open ridges, oak-laurel forest, and forest domina-

ted by Leptùspermum and üaerydiurn. Other

associâtes include Nepenthes rajah. Probably ail

populations occur vvitbin tbe boundary of

Kinabalu Parle, whose protection is therefore cru¬

cial to the continued survival of this species.

Materiai hxaminku.—Bornéo: Sabah: Barkman

& Buin 141. I.ayang I.ayang, 2600 ni, fl., 1 Mar.

1995 (*KIN); Beu ma n 9 132, Ranau District, East

Mesilau River, nr, Mesilau Cave, 6°03'N-116°36’E,

1950-2100 in, fl„ 26 Mat. 1984 (K, L); Berman

9837 , Ranau District, Pig Hill on E side of Mt.

Kinabalu, 6"03'N-1 I6"36T, 2000-2300 m, st., 25

May 1984 (A, K, L); Cbew & Corner RSBN 4364,

Mr. Kinabalu, Pig Hill, 7000 h., fl., 18 feb. 1964 (K,

L, SAN), Cbew & Corner RSNB >508, ibid., yfr., 24

Eeb. 1964 (A, K, L, SAN); Chem & Corner RSNB

4755, Mt. Kinabalu, Mesilau Cave, 6000 fr., st., 29

Mar. 1964 (K, L.. SAN); / à M.S. Clemens 50793,

Mr. Kinabalu, Gurulau spur, 7000-9000 fr., 0.. 6

Dec. 1933 (A. BM, K. L); J. & M.S. Clemens 50877,

ibid., 8000 ft . fr., 8 Dec. 1933 (A, BM, K, L);

Hopkins & Bradfbrd 5011 , Mc. Kinabalu, Pig Hill.

116°38’E-6°03'N, 2250 m, yfr., 5 Mar. 1996 (K1N„

P, MO); Meijcr SAN 28737, Ranau District, Mt.

Tambuyokan, GÜ00-7000 h., st., July 1961 (K, L,

SAN); Meijrr SAN 34617, ibid-, 6000 ft., st,. July

1961 (SAN); Mcijer SAN 48098, Mt. Kinabalu,

Ranau District, opp. Kinabalu cave, Kinabalu Nat.

Park, S000 ft.. st„ 20 Feb. 1965 (K, L, SAN); MM

SAN 4704 L Ranau Disr. Kinabalu Park, Mesilau base

camp, 5000 ft., st., June 1964 (SAN); Nais et al. SNP

3675, Kinabalu Park, Mt. Tambuyukon, 8000 ft., 0.,

16 Mar. 1991 (K1N); Nais et al. SNP4890, ibid., bet-

ween camp III and sutnmit. 11., 6 Oct. (990 (K1N);

Philips SNP 23 18. Mesilau landslide. 11., 28 May

1986 (KIN); Whithead s.n., Kinabalu, 9000 ft., st.,

Mar. 1888 (BM).

(*KIN refers to the herbarium at Kinabalu Park

HQ).

As noted in the type description, this species

appears to be most closely related to W. urdanet-

ensis from the Philippines and the Highlands of

New Guinea. In both species therc are numer-

ous, small leaflets arranged at right-angles to the

leaf rachis and the secondary veins are at right-

angles to the midrib. Weinmannia clemenisae has

characteristically bullate leaflets with the margins

recurved so rhat they can not be flattened. Tliere

is a slight tendency to bullate leaflets in W.

nrdanetensis but it is not nearly so pronounced.

In both species the upper cuticle is sometirnes

thickéned and the structure of the inflorescence

is similar. The seeds of W. clemensiae are comose

at each end, not hairy throughout as in W.

nrdanetensis.

3. Weinmannia aphanoneura Airy Shaw

Bull. Mise. Infbrm. 1940: 260 (1940); Bernardi,

Bot, Jahrb. Sysr. 83; 160 (1 964).—Type: P. W.

Richards 1716 (not 6716 as given in protologue),

Sarawak, Dulit Ridgc, open rnoss forest, ca. 1400 m,

11 Sep. 1932 (holo-. Kl, photo at KEP).

Shrub or tree 2-20 m high and up to 35 cm

dbh. Young woody stems + glabrous or adpressed

pubcrulent, soon glabrescent, older ones glabrous

with numerous lenticeis, nodes sometirnes ±

thickened and leaf scars prominent; branching

not usu.dly dichotomous. Stipules often persist¬

ent at ciistal nodes, suborbicular and usually liât,

not recurved, up to 1.7 X 1.9 cm (rarely spath ul-

ate, ca. 0.6 X 0.4), base constricted, apex broadly

rounded, abaxial surface glabrous or shortly

strigose especially towards the base, adaxlal surfa¬

ce glabrous. Leaves imparipinnate, with (0-)l-4

(-5) pairs of latéral leaflets, total length up to

16 cm, including pétiole of 0,7-2,8 cm; rachis

segments 0.5-1.9 cm long; petiole and rachis

semitercte, adaxial sidc flattened, slightly ridged

or channellcd, sometirnes narrowly winged, espe-

cially towards the point ol insertion of the Icaf-

lets, wings extending outwards up to 1 mm on

either side of raid line, petiole and rachis glab¬

rous or tomentose either on adaxial or abaxial

sutfitee but not both, usually inserted at an acute

acroscopic angle; latéral leaflets narrowly ellipt-

ical or narrowly obovate, the largest 2.4-7 X 0.6-

1.9 cm, the base unequal, ± sessiie, cuneate, apex

acute to obtuse; apical leaflet narrowly elliprical,

narrowly obovate to obovate, scareely larger than

the largest latéral leaflets, 3-9.4 X 0.7-2.7 cm

including the narrowly cuneate to attenuate base

0.3-1.2 cm long, apex acute to obtuse; blades

coriaceous, glabrous and sometirnes shiny above,

glabrous below, characteristically drying grey or

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

Fig. 10.—Distribution of Weinmannia aphanoneura, W. clemensiae and W. hooglandii.

dark chestnut above and chestnut below; not

punctate; margin crenate, 6-11 norches on each

side ol thc largest latéral leaders; midrib slightly

depressed above, prominent below and some-

times pubescent to tojnenrose at base, secondary

and tertiary venarion ± fiat un both surfaces or

often obscured especiallv on the upper surface by

the thick cuticle; réticulum not dense.

Inflorescence 1 or 2 pairs of opposite dyads, the

apical bud of the main stent between the central

dyads shortly sericeous, rarely continuing to

grow vcgctatively during floweting; bud at apex

of peduncle in angle between racemes shortly

sericeous; sometimes dyads produced in axils of

leaves at subdistal nodes; peduncles ca. 0.3-

1.9 cm long, racemes usually not longer than the

leaves (7-8 cm) but sometimes up to 14 cm long;

peduncles glabrous or adpressed puberulenc, axes

of racemes puberulent, rarely tomentose; stipules

at node at top of peduncle sometimes persistent

and either free and orbicular or connare at base

to fornt a cup-shaped structure. Floral buds

inserted in fascicles; floral bracts carinate, ca.

1.6 mm long, margin with minute glandular

tceth, shortly hairy on abaxial surface. Flowers

unisexüal; pedicel 1.8-3.6 mm long, pubcrulous;

calyx lobes 0,5-0.8 x 0.4-0.9 mm, glabrous;

corolla often rotate, petals oblong or irregularly

obovate, 1,2-1.5 x 0.7-0.8 mm, rounded or

emarginate at apex; dise lobes 0.3-0.5 mm long,

oblong and discrète or with thin flanges on

either side so thar the 8 lobes fornt an alrnost

continuons dise; in male flowers: filaments 2.1-

2.6 mm long, ovary 0.5-0.7 mm long, pubes¬

cent, styles 0.1-0.2 mm long, incurved; in feinale

flowers; filaments up to 1.7 mm long, ovary' ca.

1 mm long, densely pubescent, styles ca. 1.8 mm

long, straight, stigmas capitale, papillose.

ADANSONIA. sér. 3 • 1998 • 20(1)

Weinmannia in western Malesia

Capsule 2,5-3.5 X 1.3-1.5 mm at dehiscence,

the styles up to 15 mm long, che exocarp pubes-

cent; calyx lobes persistent; central column

sometimes présent. Seeds ca. 0.9 mm long,

comose at both ends, the hairs to 1.5 mm

long.—Fig. 9A-G, 10.

BREED1NG SYSTEM.—Dioecious.

FlELD CHARACTERS.—Bark scaly dark brown,

outer bark reddish, inner bark brownish, sap

wood pale yellow (Pitty dr Ogata SAN 63292).

Stem pink; pedoles, rnidrib on underside of leaf-

lets and inflorescence sometimes magenta. Buds

red; flowers white to pink; calyx pink or red;

corolla cream, pale pink or reddish; filaments

white with pale pink anthers. Fruits crimson,

Distribution and ecoloüy.—M ontané fcrest

in Bornéo and Sumatra (one record), at 1175-

2560 m. Habitats desuibed as open moss foresr,

ridge top forest, submontane heath lorest and

ericaceous mossy fotest. Locally comrnon.

MATERIAl 1 XAMINM 1 .—SUMATRA: Nagamasu 3641,

West Sumatra, sunimit of Gn, Rasam, ca. 10 1cm S of

Danau Diatas, 1 “ 1 0-11 ’S-l00 D 44-45’E. 2500-

2565 m. fl.. 18 Feb. 1989 (I.).^Bornf.O: Sabah:

Aban SAN 76507, Mt. Kinabalu, Ranau District,

above power station, st., 11 Mar. 1973 (SAN); J. &

M.S. (Siemens 29476, Mt. Kinabalu, fenompok, 5000

ft„ fl.. 2 May 1932 (A, BM, K, L); J. à MS Clemens

33076, Mt. Kinabalu. Marai Parai, beadwaters of

Sadikan, 5000 ft., fl., yfr, 5 May 1933 (A, BM, K, L).

Sara wAK: Anderson 4507, Baram District, Gn. Mulu,

path from Melinau Paku, 4800 ft.. buds, 30 June

1961 (A, K, 1,); Rrunig S 8785, Binrulu District,

Merurong plateau, Bt. Bandong, fl,, 15 May 1960 (A,

K, L); Burtt & Martin. 5487, 4th Div., Gn. Murud,

3°55 N-l 15 U 31'E, 7800 ft.. fl., 12 Oct. 1967 (SAR);

Chew Wee-I.ek 380 , Baram District, Gn. Mulu, 4°

5'N-114”55'E, 5000 ft., buds, 16 June 1962 (A, K,

L); Chew Wee-Lek 388, ibid., fl., 16 June 1962 (A, lv,

L); Mas Paie S26481 , Gn. Murut, Ba Kelâlan, Lawas,

7700 ft., fl., 10 Ocr. 1967 (K, L); Mas Paie S 26531,

Gn. Murut, Ulu Pahdapat, Ba KclaJan, l.awas, 7750

ft., fl., 13 Oct. 1967 (K, L). Kalimantan: Endcrt

4125, Ond. Afd. W. Koetai, b/d Kcmoel, 1500 m, fr.,

14 Oct. 1925 (U.

W. cf. aphaiumaneura. BORNEO: SABAH: Pitty &

Ogata S 63292, Tenom District, Gn. Anginon

Sapong, 3900 ft., fl., 20 Nov. 1968 (A); Ogata 11614,

ibid., 1175 m, 20 Nov. 1968 (L).

Local NAME.—Maba (Iban).

4. Weinmannia hooglandii H.C. Hopkins

& J.C. Bradford, sp. nov.

A W. fraxinea folioiis lateralibus parvinribus (maximis

< 2 tvi lottgis, mm > (2,8-) 3.5 on longis) ablongis apice

late acuto (non lanceoiatis apice auguste acuta), venatio-

ne reticu/ata tvnferta. area lis parvinribus. inflorescentia

unica pare dy aluni lateralium (non 2 net 3 paribus dya-

tum vel tetratum) diversa.

Type.— T.C. Wbitmore FRI 12582, Peninsular

Malaysia, Pahang/Sclangor, Gn. Ulu Kali, foresîed

gully, 5600 ft., 9 Sep. 1969 (holo-, KEP!; iso-, A!, K!,

I !. SAN!).

Shrub or small tree 1.2-5 m X 4 cm dbh. Young

woody stems tomentose or velutinous, older ones

glabrescent with nu mérous lenticels, branching

not dichotomous. Stipules persistent or nor at

distal nodes, ± orbicular, up co 0.5 X 0.6 cm,

base constricted, apex broadly rounded or flat-

tened, abaxial surface shordy strigose especially

towards the base, adaxial surface glabrous. Leaves

imparipinnate, with 3-8 pairs of latéral leaflets,

total length 5-7 cm long; pétiole 0.8-1.5 cm

long, rachis segments 0.4-0.8 cm long, pedole

and rachis terete, slightly wider towards the point

of insertion of the leaflets, otren densely toment¬

ose and especially so along adaxial s idc; latéral

leaflets ± oblong to narrowly ovate, slightly

imbricate the Jargest 1.6-2 X 0.5-0.6 cm, the

most proximal leaflets smaller, ca. 0.7 X 0.4 cm,

base unequal, sessile, cuneate, apex broadly

acute; apical leaflet narrowly elliptical, scarcely

lurger than the largest latéral leaflets, 1.5-2. 3 X

0.5-0.8 cm, base attenuate, apex acute; hlades

coriacenus, glabrous and sometimes sliiny above,

glabrous below, not punctate; margin crenate, ca.

3-7 notches on each side of the largest latéral leaf¬

lets, margin minutely revolute; rnidrib slightly

depressed above, prominent below and some¬

times shordy strigose especially towards the base,

secondaty and tertiary venation i fiat on both

Surfaces or often obscure on the upper surface,

réticulum relatively dense and drying darker

than the remainder of the blade.

Inflorescence a pair of latéral dyads, the apical

bud of main stem between the peduncles shordy

sericeous; bud at apex of pedunde between

racemes shortly sericeous; peduncles ca. 0.4-

0.5 cm long, strigose; racemes up to 6 cm long.

ADANSONIA, sér, 3 • 1998 • 20(1)

Hopkins H.C.F.

the axis pubescem or tomenrose. Floral buds

inserted in fascicles; llora! bracts carinate, ca.

1 mm long, minurely hairy; pedicel 1.5-3 mm

long, minurely hairy. Male flowersi calyx lobes

ca. 0.8-0,9 X 0.6 mm, glabrous; petals ± obovate,

1.2-1.4 X 0.8-0.9 mm, apex roundcd or emarg-

inate, minurely ciliolate (atx 40); dise lobes 0.3-

0.4 long, oblong or broadly oblong wich thin

flanges on either side of each lobe; filaments ca.

2.6-3 mm long; ovarv minute, ca. 0.6 mm long,

densely golden pubescent, styles 0,1-0.2 mm long,

incurved, Fruit unknown.—Fig. 10, 11A-D.

JUVENILE FOLlACîE.—Non-reproductive (copp-

ice ?) shoots; axes tomenrose, internodes long,

often ca. 8 cm. Stipules orbicular to reniform

and shortly sralked, up to 1 X 1.2 cm. Leaves up

to 11 cm long, almost perpendicular to stem

(not ± erect as in adult foliage), latéral leaflets in

8-10 pairs, ± oblong, up to 2 X 0.8(-l) cm,

chartaceous sometimes widely spaced on the

rachis (not + imbricate); rachis segments up to

0.9 cm. (See Hoogltind 1257S, Si eu,' Wct Hoe 15

and Chiew Geôle Juvénile leaves are longer

than adult ones and hâve moce pairs of leaflets.

BreEDING SYSTEM. —Probably dioecious, but

fentale flowers hâve not been seen.

FlEl.D CHARACTËRS.—The leaves are clustered

towards the end of the twigs and held ± erect to

form an even, dense, ball-like, congested crown

(J.C. BlUUHORD pers. obs ). Young branches and

leaf rachises purplish or brown, Foliage dark

green above, leathery. Flowers wliire on pink

stalks ( Wbitmore FRI 12582), corolla and sta-

mens white (Ng FRI 6236).

Distribution and ecology. —Known from

mountains in western Pemnsular Malaysia, from

ca. 1680-2065 m. Habitats described as upper

montane fbrest, moss forest and summit forest of

low stature; locally common.

Fig. 11.— Weinmannia hooglandii A, Flowerlng shoot showing inflorescence of two “dyads" (each of two racemes on a short

peduncle) in opposite leaf axils al lhe mosl distal node of the shoot; B, Underside of a latéral leaflet; C, Node on a végétative shoot

showing a pair of stipules between opposite pétioles with a sericeous bud in the axil of the right-hand petiole; D, Male flower, the fila¬

ments not fully extended. (A, B, D, Whitmore FRI 12582-, C, Bradford 580 ).—Drawn by C. Monnié and D. Storez.

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in western Malesia

Material examiner. —Peninsular Malaysia:

Bradford 579, 580, Pahang. Gn. UJu Kali, Genting

Highlands, 3°23’N-H)r45'E, 1680 m, st., 18 Mar.

1996 (KEP, MO, P); Chiew Gcok s.n., Pahang,

Genting Highlands, 5600 ft., st., 22 J une 1974

(BISH); Hoogland 12578 , Pahang, Sutntnit Gn.

Brinchang, 6300 )t.. st., 1 Apr. 1975 (CANB);

Murdoch s.ii,, Selangor, Gn. U lu Kaii, st., 25 Mar.

1905 (K): IVg FRI 6256, Selangor, Summir Gn.

Mengkuang, il, 1 ! Jan. 1972 (KEP); Stew Wei Hoc

15, Gn. Ulu Kali, Gcnring Highlands. 5800 fr., st., 4

June 1977 (E); Soevadmo Ç r Suh.umi $ 172, Gn. Ulu

Kali, 3°27’N-103“48’E, 1772 m, st., 8 May 1990 (A);

Symingion 32246 , Perak, Gn. Korbu P.R., Camcron

Highlands. 5000-7000 fr., st.. 22 July 1933 (KEP).

This species is named for R.D. HOOGLAND,

specialist in rhe Cunoniaceac (or over 35 years,

who recognized his collection from Gunung

Brinchang as “something interesting”.

Rel AT IO N S HI PS.—The material placed here in

Weinmannia hooglandii had previously been

regarded as a small-leaflcted variant of the com-

mon and widespread W. fraxinea (D. Don) Miq.

(synonym W. blumti Planch.) from high éléva¬

tions. However, J.C. BRAWORl) observed indivi-

duals of VP! hooglandii growing nexr to W

fraxinea (with much larger, narrowly elliptical

leaflets) at the same altitude and in the same

habitat at Gn. Ulu Kali, Genting Highlands, in

March 1996, suggêsting that W. hooglandii is not

merely an ecotype of the latter, and could be

regarded as distinct. This observation is docu-

mented by Bradford 578 and 581 (W. fraxinea)

and Bradford 579 and 580 ( W. hooglandii).

Weinmannia hooglandii is distinguished from

W. fraxinea by its srnaller leaflets with more den-

sely reticulate venation visible on the lower surf¬

ace, and shorter inrernodes producing a

congested, not diffuse, crown. The shape of the

latéral leaflets is also different; in U7 fraxinea in

Peninsular Malaysia, the leaflets are usually lan-

ceolate or narrowly elliptical (lengtli to breadth

ratio 1: 0.23-0,33(-0,38)] while in W hooglandii

distal leaflets are oblong or subfalcate (with the

main vein asymetrically placed) or sometimes

suboblanceol.rre (length to breadth ratio 1: 0.30-

0.39) and the proximal ones broadly elliptical. In

addition, the stipules m W hooglandii are small-

er, being up to 6 mm broad, while the smallest

seen in U’.' fraxinea are 9 mm broad, though in

many specimens they are not seen, having alrea-

dy fallen,

A further différence between Weinmannia hoog¬

landii and W. fraxinea is the presence of super-

numerary latéral buds at nodes, especially those

bearing partial inflorescences, in the latter.

Usually in W fraxinea a reproductive node bas

two or three buds in each opposite leat axil, and

one, two or three pairs of partial inflorescences

develop from them (each partial inflorescence

being a dyad or a terrad). In the few fertile speci¬

mens of W, hooglandii available, there is only one

pair of buds ar a reproductive node, and so the

inflorescence that develops consists of a pair of

dyads (Eig. IIA), as in W. urdanetensis (see

below).

AJthough Weinmannia hooglandii and W fraxin¬

ea can readilv bc distinguished from one another

where they occur together, W. fraxinea is a am¬

ples and variable species that in some parts of its

range has rather small leaves whose leaflets

approach those of W. hooglandii in size [e.g.

Cockburn FRI 11030, Peninsular Malaysia,

Pahang, Summit Padang of Gu, Tapis, 4500 ft.,

fr., 11 Sep. 1968 (K, KEP, l ); Symington & Kiah

SF28894, Pahang, Gn Tapis, Kuantan, 460U ft.,

fl., fr., 14 June 1934 (BO, SIN G ) ]. In these

cases, leaflct shape and the venation are typical of

W. fraxinea, and the leaflets are more widely

spaced (not imbricatc) along the rachis.

Weinmannia urdanetensis, which grows in mon-

tane forests in New Guinca and the Philippines,

also has small leaflets, and like W hooglandii, the

inflorescence is rypicaüy a pair of dyads. Mature

foliage of W. hooglandii differs from thar of W.

urdanetensis by its usually shorter leaves with

fewer pairs of leaflets [4-8 not (2-)6-10(-19)], the

leaves held ar an acute angle to the stem in mat¬

ure loliage (not more or less at right angles) and

the leaflets positioned at an acute angle to the

leaf rachis (nor more or less at right angles).

Acknowledgements

The work by HCFH was fmanced by the Human

Capital and Mobility Programme of the European

Union via the network on Botanical Diversity ot the

Indo-Pacific. The following herbaria are thanked for

access to specimens: A, B, BISH, BM, BO, CANB,

AbANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

GH, K, KEP, L, LINN, MA, NOU. P, S, SAN, SAR,

SING, SUVA. US, Z. Jason Braphord is gratefuJ for

financial support from Washington Univcrsïry, indud-

ing the NSF Graduate Research Training Support

Grant NSF BIR-9256779, and from the Missouri

Botanical Gstrden, espetially tlirough the Mellon

Foundation Fellowsliip. For assistance in our field-

work in 1996, we are greatly indebted to John and

Tefîÿ BeaMAN (UN1MAS, Sarawak), Jamili Nais and

his staft (Kinahalu Park, SâbahJ, S.AW LliNG CUAN

and his staft (FR1M, Peninsular Malaysia), Michael

Doyge (SUVA, Fiji) and the Fiji Electricirv Authoriry

(Monasavu Dam), the Directors of the ‘ Service de

l’Environnement" fot the North and South Provinces

in New Caledonia, and Tanguy Jaffré and his staff at

ORSTOM in Nouméa. HCFH also rhanks the late

R.D. HouGEAND for his generosity in giving access to

his unpublished notes, Jason Bradpord for nitich

interesting discussion on thosc parts of the text for

which he is not cired as co-author, and lot the pboto-

graphs fot Fig. 2, Petra HofFMANN for advice on

nomenclature, P. Lo'X'RY & G. MCPHERSON for cotn-

ments on ail or part of the manuscript, Eisa AR.MÈDI

for help wirh typing, and Cécile MONNtf, Dominique

STOREZ and Jacqueline LEMEl'X for the illustrations.

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ADANSONIA, sér. 3 • 1998 • 20(1)

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ADANSONIA, sér. 3 • 1998 • 20(1)

A révision of Weinmannia (Cunoniaceae)

in Malesia and the Pacific. 2. Sulawesi and

the Philippines

Helen C.F. HOPKINS

Laboratoire de Phanérogamie, Muséum national d'Histoire naturelle,

16 rue Buffon, 75005 Paris, France.

Masons Arms, Hutton Roof, via Carnforth, Lancashire, LA6 2PE, U.K.

KEYWORDS

Weinmannia,

Cunoniaceae,

Sulawesi,

Philippines.

ABSTRACT

Eleven species of Weinmannia ottur in Sulawesi and ihc Philippines, but

only one (W. negrosensis) is commun to both régions. Ail species are placed in

sect. Fasciatiata although W. descornbesiana shows some charactcristics of

sect. Leiospemum. Of the seven specics from Sulawesi, four are described

here as new: W. dcvogelii is closely related to W. fiaxinea, the most wîde-

spread Malesiati species which is absent from both Sulawesi and the

Philippines; W. furjuracea also occurs in Seratn; W. caadei and W. eyrnaeana

are both endémie. The seventh species from Sulawesi, W, célébrât, is rather

poorly known. In the Philippines, five species are recognised, noue of them

new. Weinmannia huicbinsonii, W. luzoniensis and W. lucida are ail endemic,

W. urdanetensis occurs in both the Philippines and New Guinea, and the

fifth species is W. negrosensis, Keys, illustrations and distribution maps are

given.

MOTS CLÉS

Weinmannia,

Cunoniaceae,

Célèbes,

Philippines.

RÉSUMÉ

Onze espèces de Weinmannia existent aux Célèbes et aux Philippines, mais

une seule (W. negrosensis) est commune aux deux régions. Toutes les espèces

appartiennent à la sect. Fa si ieu la ta, bien que W, descornbesiana présente cer¬

taines affinités avec la sect. Leiospermunr. Parmi les sept espèces des Célèbes,

quatre sont nouvelles et décrites ici : W. deoogelii, très affilié de W, fmxinea

qui est l'espèce la plus répandue en Malésie mais absente des Célèbes et des

Philippines ; W, furjuracea qui existe aussi à Sent ni ; W. eoadei et W, eyrnaea¬

na, toutes deux endémiques, lut septième espèce, des Célèbes, U 7 , eelebica, est

assez peu connue. Cinq espèces existent aux Philippines, mais aucune n’est

nouvelle. Weinmannia hutebinsonii, W. luzoniensis et W. lucida sont endé¬

miques, W. urdanetvnsts sc trouve à la fois aux Philippines et en Nouvelle-

Guinée, la cinquième espèce étant W. negrosensis. Des clés de détermination,

des illustrations et des cartes de distribution sont présentées.

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

This is the second part of a révision of the

Malesian-Pacific species of Weinmannia. Part 1

(HOPKINS 1998a) gives a general introduction,

including définitions of the terms used for the

structure of the inflorescence, and describes the

species of western Malesia. Parts 3 and 4

(Hopkins 1998b; Hopkins & Florence 1998)

deal with the species of the western and central

Pacific respectively.

Most of the Weinmannia species found in

Sulawesi and the Philippines are clearly defined

and monomorphic. Most are also endemic to

one or other of these régions, though there is a

marked différence in that ail the taxa recognised

here from the Philippines hâve been known for

some time, while for Sulawesi, four species are

described here as new, and incomplète material

probably represents a flfth. Sulawesi has the

highest number of sympatric Weinmannia species

of any island (or island group) in Malesia, and

further exploration of its montant forests might

well be rewardmg. Although ail the species dealt

with here be|ong to sect. Fascieulata. W. descom-

besiana from Sulawesi is noteworthy because it

shows some characteristics of sect. Leiospermum

(see below).

WEINMANNIA IN SULAWESI

Seven species arc recognised from Sulawesi

(Celebes), five of which are endemic

( Weinmannia cdebica > W. coodei , W. descombes-

iana , W devogelii and W eymaeana). AU occur in

montant forest with the exception of W. devogel-

ii , which occurs in lowland forest up to 700 m.

Two species hâve bisexual flowers ( W. descombes-

iana and W. furfuracea), four hâve unisexual

ones and for W coodei , the breeding System is

undetermined.

1 .

2 .

2 ’.

3’.

4’.

5’.

6 .

6 ’.

Key to the species in Sulawesi

Leaves simple (unifoliolate); flowers inserted singly in the axils of the bracts on the inflorescence axes .

..... 1. W. descombesiana

Leaves trifoliolate or imparipinnate (or rarely unifoliolate); flowers inserted in fascicles in the axils of the

bracts on the inflorescence axes ...2

Leaves 1-3 foliolate, coriaccous ....7. W. negrosensis

Leaves usually with 3 or more leaflets, charraceous to subcoriaceous .3

Latéral leaflets in 3-20 pairs, the largest per leaf 0.7-1.9 X 0.2-0.5 cm .4. W. eymaeana

Latéral leaflets ni (0-) 1-6 pairs, the largest per leaf & 3x1 cm . 4

Latéral leaflets elliptical to avare, apex acuminarc.5

Latéral leaflets narrowly elliptical to obovate, apex obtuse or rounded .6

Pedicels 0.2-0.5 mm long at anthesis; latéral leaflets 4.5-10 X 1-3 cm; indumenrum on young stems sparse

and never pilose .2. W, devogelii

Pedicels 1.8-2 mm long at anthesis; latéral leaflets 3-5 x 1-1.6 cm; indumentum on young stems at least

pardy pilose .....6. W. celebica

Young stems and leaves, stipules, leaf rachises and inflorescence axes bearing a rnsty or golden brown vill-

ous-tomentose indumentum .........3. W. furfuracea

Young stems glabrous; leaf rachises glabrous or comentose on adaxiaJ side, inflorescence axes puberuious;

indumentum neither rusty nor golden brown ...5. W. coodei

1. Weinmannia descombesiana Bernardi

Bot. Jahrb. Syst. 83: 190, t. 33 (1964).—Type:

Kjellberg 1618 , SW Sulawesi, Pasoei-Rante Lemo,

1000 m, fl., 9 June 1929 (holo-, S!; iso-, BO).

Shrub or tree, 2.6-30 m high, up to 30 cm dbh.

Young stems glabrous or puberuious, black,

shiny, older ones with numerous white lenticels;

branching sometimes dichotomous. Stipules

usually caducous, obovate, ligulare or spathulate,

ca. 0.7 X 0.4 cm, apex rounded, abaxial surface

shorrly strigose especially towards rhe base,

otherwise glabrous, adaxial surface glabrous.

Leaves unifoliolare usually with an articulation

between blade and petiole: petiole 0.5-1 cm

long, semiterete and somewhat channelled, rarely

winged, almost glabrous; leaf blades elliptical.

ADANSONIA, sér. 3 • 1938 - 20(1)

Weinmannia in Sulawesi and Philippines

\ 4 5 mm

V \ -

I J

{Sit

il f

c. rtoMWté

Fig. 1.—A-F. Weinmannia devogelii. A, flowering shoot with one of a pair of imparipinnate leaves and an inflorescence of 4 pairs of

tetrads; note minute Duds in angle Between the central racemes of each tetrad; B, male flower, styles too short to be visible; C,

female flower with very short filaments; D, dehlsced capsule; E, seed; F, stipule. (A, B, Meijer 11147; C, de Vogel 5682 ; D-F, de

Vogel 5959). —G-L, Weinmannia descombesiana G, shoot showing infructescence developed from an axillary shoot bearing 3

dyads, the apical bud continuing to grow vegetatively; H, section of axis of a raceme showing pedicels of capsules inserted singly; J,

bisexual flower; K. dehisced capsule; L, seed. (G, H, J. bb2087ù, K, L, bb 22981).— Drawn by C. Monniè.

ADANSONIA, séf. 3 • 1998 • 20(1)

Hopkins H.C.F.

4-9.5(-11) X L7-4.4(-4.8) cm, base acute, apex

acuminare, subcoriaceous or cbartaceous, glab-

rous on borh surfaces, not punctate below; mar-

gin sometimes minutely thickened and revolute,

crenate, ca. 8-10 crenations on eacb side; tnidrib

flat or slightly depressed above, glabrous, promi¬

nent below with sparse indumentum, secondary

and tertiary vcnation fiat above and below, réti¬

culum dense.

Inflorescence usually a pair of dyads', sometimes

a few successive nodes producing partial infloresc¬

ences simultaneously; peduncles 0.2-1.6 cm

long, sparsely puberulent; when inflorescence is

at most distal node of stem, apical bud betwecn

the central pair of peduncles usually présent and

sericeous or rare!y developed into a partial inflor¬

escence; buds at apex of peduncie in angle bet-

ween central pair of racemes sericeous, minute;

axes of racemes puberulent, up to ca. 8.5 cm

long. Floral buds inserted singly; floral bracts

carinate, ca. 0.7 mm long, strigose-pubescent.

Flowers bisexual; pedicel 1-1.2 mm long in flow-

er, 1.4-2.6 mm long in fruit, alrnost glabrous;

calyx lobes 0.5 X 0.4 mm, sparsely hairy on outer

surface, dilate; petals oblong, apex roundcd, 1.2

X 0.6 mm, minutely ciliate; dise lobes 0.2-

0.3 mm long, oblong, minutely strigose; fila¬

ments ca. 2.2 mm long; ovary ca. 1 mm long,

densely pubexcent; styles ca. 1 mm long, diverg-

ing; stigmas capitale, papillose.

Capsule 2-2.5 X 1.2-1.6 mm at dehiscence, the

styles ca. 1 mm long, the exocarp strigose-

pubescent; calyx lobes persistent; central column

présent and as long as valves. Seeds 0.5-0.6 mm

long, ca. 8 per capsule, often persisting in cap¬

sules after dehiscence, comose at boih ends, the

hairs to 1.5 mm long, hairs alrnost “sticky”.—

Fig. 1G-L, 2.

JUVENILE FOLIACE. —Stipules suborbicular, not

amplexicaul, up to 1.5 X 1.3 cm, persistent.

Leaves elliptical, the petiole up to 1 cm long, the

blade up to 17.5 X 6.3 cm, chartaceous, the mar-

gin with up to 20 nocches on cach side.

BRF.EDING SYSTEM. —Hcrmaphroditic.

FlELD CHARACTERS. —Young leaves red or redd-

ish purple. Buds pink or green, flowers whire or

pale green, smell faint; fruits yellow or green,

tuming red with âge.

Distribution and ecology. —Mountains of

South and Centrai Sulawesi, at 1000-1800 m. In

primary and secondary forest on a variety of soil

types.

Mau RIAL EXAMINEE).—SOUTH SutAWESI: bb

20870 , Ond. afd. Mamasa, nabij l andoeng, 1400 m,

buds, fl., fr., 13 July 1936 (A, K, L); bb 2298], Ond.

afd. M.ikule-Ratepao, nabij Tandoeng, 1150 m, fr.,

17 July 1937 (A, BUSH, BM, L, P); bb 24206, Ond.

afd, Mamasa, nabij Limbong, 1500 m, yfr., 7 Apr.

19.38 (A, L); b b 26643, Ond. afd. Masamba, nabij

Tcdcboe, 1800 m, fr., 25 Nov. 1938 (A, L); Mamakit

bb s.n., Ond. afd. Mamasa. nabij Mamasa, 1150 m,

sr., 2 Aug. 1941 (F); F.yrrut 432, Ond. ifid. F.nrekang,

betwecn Pasoci-Rante Lemo, 1330 m, st., juv., l4

(une 1937 (L); Fyma 3590, Rec. Menado, Ond. afd.

Poso, between Biv. III - Biv. IV. top Gn. Loentoet,

IV., 4 Sep. 1938 (A, K, L).

LOCAL NAMES, —Paseh (language Toradja, dialect

Tai), Panessean (Passe) (language Toradja),

Londong (languages Toradja & Rongkong),

Malètoea (language Ram pi). Local uses: firewood.

This species was placed in secc. Leiosperrnum

{sub nom. sect. Racêmosue) by BERNARD! (1964)

because the flowers are arranged singly in the

axils of the bracts on the axes of the racemes.

However, it shows a mixture of characters of

borh sect. Leiosperrnum and Fusciculara as they

are circumscribed in part 1 (Hopkins 1998a).

Besides the insertion of the flowers, those charac¬

ters seen in Wêinmannia dtseombesmna that are

indicative of sect. Leiosperrnum are; branching

sometimes dichotomous; stipules ovatc-ligulate-

spathulate in adult foliage; free central column of

capsules well developed.

However, the inflorescence structure is ty'pical

of sect. F&sciculata, and usually consiscs of a pair

of dyads or tetrads with short, laterally ttattened

peduncles, that develop from axillary buds.

Whcn the dyads are at the most distal végétative

node of a shoot, rhere is usually an apical bud

between them or rarely it may develop ijlto a

thïrd dyad (sec Fig. 1 G); in some other cases the

partial inflorescences are inserted in the axils of

leaves at subdistal nodes. Other characters that

suggest an affinity with sect Fasciculata are the

pubescent (not strigose) indumentum on the

capsules, the calyx lobes persistent in fruit, and

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

the absence of “collars”, which are ihe remnants

of a pair ofpartially fused, protective stipules, on

the axes of the partial inflorescences, and which

are seen frequently in sect. Lnospcrmum. A clad-

istic analysis based on morphology suggests that

this species belongs in sect. Fasciculata

(BRADFORD in press).

2. Weinmannia devogelii H.C. Hopkins,

sp. nov.

A Weinmannia fraxinea (D. Don) Miq. floribus sub-

sessilibus (pedicellis 0.2-0 .5 mm loti vis non ( 1.1-)!.5-

3 mm) attingentibus roseis non albldis diffèrt.

Type. — de Vogel 6122, (South Sulawesi) Sulawesi

Selatan, N shore of Lake Matano, Bonemaitu, E of

Nuha, 2°20’S-121°27’E, 400 m. fl., 5 July 1979

(holo-, LT; iso-, K!).

Tree 5-20 rn, up to 40 cm dbh. Young stems

woody with sparse indumentutn, older ones glab-

rescent with minute longitudinal fissures and

numerous lenticels; branching not dichotomous.

Stipules caducous or not, ± orbicular or larger

ones amplexicaul, up to 1.4 X 1.6 cm, base

usually constricted, apex rounded, adaxial surface

giabrous, scurfy or puberulent, abaxial surface

shortly strigose or glabrescent. Leaves imparipinn-

ate, latéral lcaflets (0-)2-6 pairs, total length up

to 22 cm; pétiole 1.5-3-5 cm long, rachis seg¬

ments 1.2-2.5 cm long, petiole and rachis seg¬

ments terete to semiterete, somewhat flattened

and densely pubcscent on adaxial side, indument-

um sparse on abaxial side; latéral leaflets lanceo-

late, narrowly ovate or narrowly elliptical, the

largest 4.5-10 X 1-3 cm, the base unequal, ± sess-

ile or shortly petiolulate, apex acuminate; apical

leaflet narrowly elliptical, ca. 5.5-9.5 X 1.5-2 cm,

base attenuate into a petiolule ca. 0.7 cm long,

apex acuminate; leaflet blades chartaceous to

subcoriaceous, giabrous above and below, not

punctate; margin alrnost entire to crenulate, 10-

15 notches on each side of the largest latéral leaf¬

lets; midrib slightly depressed above, puberulent

at base, prominent below and sparsely strigose,

secondary and tertiary venation slightly raised on

both surfaces, réticulum not dense.

Inflorescence of 1-3 pairs of dyads or tetrads.

sometimes a fevv successive nodes producing part¬

ial inflorescences simultaneously, pedunclcs latér¬

al ly flattened, 0.2-1.3 cm long, puberulous;

bud(s) between the central pair of peduncles scric-

eous; buds at apex of peduncle in angle between

central pair of racemes sericeous, minute; axes of

racemes densely puberulous, up to 11.5 cm long.

Floral buds inserted in fascides; floral bracts ±

carinate, ca. 0.8 mm long, minutely strigose,

caducous, Flowers unîsexual; pedicel 0.2-0.5 mm

long, minutely strigose or puberulous; calyx

lobes 0.5-0.7 x 0,4-0,5 mm, minutely strigose

on outer surface; corolla obovate, 1,1-1,5 X 0.7-

1.1 mm, apex rounded or irregularly entarginate,

ciliate; dise lobes 0.2-0,3 mm long, oblong; in

male flowers. filaments 3.3-3.7 mm long, ovary

0.4-0.5 mm long, pubescent, styles 0.1-0.2 mm

long, incurwed; in fernale flpwers: filaments 0.6-

0.9 mm long, ovary 0.9-1.1 mm long, densely

pubescent. styles 0.9-1.1 mm long, straight, stig¬

mas papillose,

Capsule 1.9-2,5 x 1.2-1,5 mm at dehiscence,

the styles up to 1 mm long, the exocarp pubesc¬

ent; calyx lobes persistent; central column prés¬

ent. Seeds ellipsoid to broadly ellipsoid,

0.5-0.7 mnt long, 4 pet capsule, comose at both

ends, the hairs to 2 mm long.—Fig. 1A-F, 2.

BRP.F.DING SYSTEM.—Dioecious.

FlEI.D CHARACTERS.—Buttresses few or absent.

Outer bark ochre-grey, fissured or not; inner

bark reddish; sapwood créant coloured; heart-

wood very hard, reddish. Young leaves red.

Flowcr buds red; flowers usually given as pink,

rarely white and fragrant ( Waturandang 18);

young fruit reddish green, old ones dark red.

Distribution and icology. —Easrem South

Sulawesi near Maliü and Lake Matano at 0.5-

700 m. In primary and secondary forest, includ-

ing disturbed primary forest with an

undergrowdi of sedges and ferns. On peridotite

and ulrrabasic latérite; also in patchcs of coastal

végétation along the lake on limestone and alluv-

ium derivcd front limestone and schist, and on

red clayey soi 1 derived from conglomerate

bedrock. Locally common.

Paratyi'es.—South SulaWESI: van Balgooy 3625,

Soroako-Wasuponda road, 2°15’-3°S, 121°-121°45’E,

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in Sulawesi and Philippines

600 m, fl., 9 July 1979 (A, K, L); van Balgooy 3809,

Soroako-Malili road nr. junction with mafl ro Larona

Hydroelectric plant, 2°1 5'S-121°45'E, 100 m, fl., 29

June 1979 (A, K, L); Damardi 2236, 2'*15’-3“S, 121'*-

121°45’E, 500 in, 11., 14 July 1979 (A, K, L); Meijer

11147, Matano Lake nr. Soroako, NE of Malili, E of

Nuha village, 2°35'S-121 U 20'E. fl.. 15 July 1976 (I.);

Meijer 11181, Matano Lake tir. Soroako, NE of

Malili, nr. old Mining office, 2°35’S-121 "20’E. yfr.,

16 July 1976 (L); Ramlanto 138, Wae Atue,

Manurung 7 km from Malili, 5 m. 11.. 10 Apr. 1984

(A, K, L); Reppie 392, CeUIII-18, Ond. afd. Malili;

nabij Oc.soc, fr., 11 Mar. 1935 (A, 130); Srhmid5512,

Soroako, 500 m, buds, |an, 1979 (E, P); de Vogel

5682, S of Lake Matano, ca. hallway herween Soroako

and Wasupondo, 2°3l'S-1 21 "21 ’E, 700 m, fl., 10

June 1979 (K, L); de Vogel 5959, S shore of Lake

Matano W uf Soroako and Taipa, on and nr. Pulau

Lintu, 2°29 S-121'15'E, a00 m, ff., 23 July 1979 (K,

L); Waturandang 18, Cei.1111-18, Ond. afd. Malili,

nabij Oesoe, 0.5 m, st., IdOct. 1931 (130, K, L).

LOCAL NAMKS. —Poémé (Tobela language, dia-

lect To Pado E).

Weinmannia devogelii is a satellite ol the wide-

spread and variable W. fraxinea. While the différ¬

ences between the twO are not greac, the first is

considered here to be more lhan a mere variety

of the second because it has a unique morpho-

logical feature (the very short pedicel) that is not

seen in W fraxinea, a distinctive ecology, and an

allopatric distribution. However, the leaf charact-

ers of W. devogelii fall within the range for W

fraxinea and stérile spccimetis catinot be disting-

uished; the latter species is assumed to be absent

from Sulawesi as no fertile material can be ident-

ified as belonging to it (unless Kjellberg 2950

(BO) [Todjamboe, 700 m, tree 10 m; flowers

male; leallets 3-5 per leafl proves to be it). The

flowers are usually pink in W. devogelii and usual-

ly white in W fraxinea at low altitude.

This spccies is named after E.F. DH VOGHL who

collccted the type. The prefix “de” is included in

the epithet on purpose

3. Weinmannia fiirfuracea H.C. Hopkins, sp.

nov.

Weinmannia devogelii H.C. Hopkins ajfrnis, sed

foliolorum indumento furfitraceo, nunquam glabra, api-

cibus juvenalibus indumento villoso-tomentoso, non bre-

viter pubescence et folialis ellipticis vel obovath apice

rotundato vel obtuso, non ovatis apice acuminato, prae-

cipuc differt.

Tvi'li.— Tantra 1595, Central Sulawesi, 0 D 30’-

1 "30’S, 119 o 30’-120°30’E, W slope Mt. Roroka

Timbu, SE of l’alu, primary forest, 2000 m, fl., 5

May 1979 (holo-, L!; iso-, BO).

Tree 16-45 m high, up to 60 cm dbh. Young

stems and leaves, stipules, leaf rachises and inflor¬

escence axes eovered with dense rusty or golden

brown villous-tomentose indumentum, the long-

est hairs up to 1 mm; older stems tontentose or

puberulous, finally glabrescent with longitudinal

fissures and numéro us lenticels, brandring not

dichotontous. Stipules caducous except at grovv-

ing tips and in immature foliage, ± orbicular,

recurved, up to ca. 0.9 X 1.1 cm, base constrict-

ed, apex rounded, abaxial surface tomentose-

villous, adaxial one velutinous. Leaves imparipinn-

ate with (l-)2-4 pairs of latéral leâflets, total

length up to 20 cm; petiole 1-4 cm long, rachis

segments 1-3.2 cm long, petiole and rachis seg¬

ments terete to semiterete at point of insertion of

leallets where sontewhat flattened on adaxial

side, villous-tomentose; latéral leâflets elliptic.il

to obovate, the largest per leaf 4-7.8 X 1.9-3 cm,

rhe base almost equal, + sessile or shortly petiol-

ul ire, apex rounded or obtuse (acute in immatu¬

re foliage); apical leaflet cllipticâl to obovate, 5-9

(-11) x 1,9-3.6(-4.8) cm including base attenu-

atc into a petiolule ca. 1 cm long, apex rounded

or obtuse; leaflet blades subcoriaceous, glabrous

above, with an indumentum of short, scurfy

hairs below, sometimes glabrescent, not punct-

ate; rnargin subentire to crenulate, 20-25 notches

on each side of the latgest latéral leallets;

midrib ± fiat above, prominent below and bear-

ing villous to scurfy hairs, gecondary and terti-

ary vénation ± fiat above and somewhat raised

below, réticulum fairly dense.

Inflorescence 1 or 2 pairs of dyads or tetrads,

the apical bud betvveen the central peduncles

often corninuing to grow vegetatively during flo-

wering, sometimes a few successive nodes pro-

ducing partial inflorescences simultaneously;

peduncles laterally flatrened towards apex, 0.6-

2.5 cm long, villous-tomentose; apical buds bet¬

ween the central pair of peduncles and buds at

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

apex of peduncles in angle between central pair

of racernes sericeous; axes of racemes tomentose,

up to 11.5 cm long. Floral buds inserted in fasc-

icles; floral b tacts not seen, Flowers bisexual;

pedicel 1-1,7 mm long, tomenrose; calyx lobes

0.7 x 0,5-0.6 mm, tomentose on outer surface;

corolla irregularly obovate to almosr circular, 1-

1.2 x 0.8-1 mm, apex rounded or irregularly

emarginate, puberulous on outer surface, margin

ciliate; dise lobes 0.3-0.4 mm long, oblong,

sometimes with thin flanges on either side to

fdrm an almost continuous dise; filaments 2.2-

2.4 mm long; ovary 1.1-1.8 mm long, densely

pubescent; styles 1.1-1.8 mm long, divergent

then later straight; stigmas capitatc, papillosc.

Capsule 3-4 x 1.5 mm before deliiscencc, the

styles up to 1.5 mm long, the exocarp densely

pubescent; calyx lobes persistent. Immature seeds

0.6-0.8 mm long, comose at both ends.—Fig. 2,

3A-D.

Brefdjnc; SYSTEM.— Fiermaphroditic.

Fiei.D CHARACTF.rts.— Tree without buttresses.

Living bark dark brown, wood yellowish, heart

wood light brown. Leaves golden from below,

young leaves brown, old ones glaucous below.

Flower buds greenish whire, flowers white, Irag-

rant, lull ot bees ( Turnra 1595). Immature fruits

yellowish green.

Distribution and ecolocy. —Mountains of

Sulawesi and Seram. Primary forest at 1700-

2000 m, including montane forest dominated by

Agathis (up to 40 m tall) and in forest dominated

by Castanopsis with Trimenia, Eugenia,

Phyllocladus , Ericaceae and Pandanus. L.ocally

common.

Paratyi’es.—Central SuLAwr.st: van Balgoay

3255 , ü“30’-1 "30'S, U9 a 30'-120°30'E, Mt. Roroka

Timbu, W slope, 2000 m, fl., 10 May 1979 (A, L);

van Balgoay 3464, OMO’-IDOS, 119°30’-l20 o 30’E,

Danau Tambing, 1700 m. Ir., 23 May 1979 (A, L);

Tantra 1587, W slope Mt Roroka Timbu, SE of

Palu, 0 e 30'-l Q 30’S, II9°30'-120 ,, 30 , E, 2000 m, st.,

juv„ 5 May 1979 (1.); Zi] U de Joug II4/V2 , bb 20787.

Ond. ald. Fnrekang, Sawito, 1000 m, fl., 14 June

1936 (BO), —Seram: Rutten 2231, Mid Seram,

Brongebergte Makina, ± 1000-1100 m, buds, 18 May

1919 (BO, L).

LOCAL NAMES. —Boeloean laki (language

Buegin, Dialect Maiwa), Ula (language Uma).

Another specimen from western Seram with

much stnaller leaflets is close to this species

(Tanah goyang, Eyma 3008 [K, L, P]) but differs

by being a large shrub rather than a tree, with

unisexual not bisexual flowers. It also resembles

W. jraxmea but that species lacks long, coarse,

dense hair.s,

4. Weinmannia eymaeana H.C. Hopkins, sp.

nov.

Weinmannia urdanetensis Elmer affinis , sedpetiolis

rbaebidibusque atnaliculatis auguste auttis et gkt bris, nec

teretibus net: dense velu finis, foliolh ellifiicis bdsi çunea-

lo non oblongis bttsi irumrtlo Vel corda to et seminibus

comosis utrintfue , non hirsutis ttbiqne, ptaecipue differt.

Tvit.— Eyma3578, Central Sulawesi, Res. Menado;

Ond. ald. Pnso, Tusschén, Biv. 11-111, op N uidooper

van Gn. Loemoet, fl., fr., 3 Sep. 1938 (holo-, L!; iso-,

Al, BO!, K!, U).

Shrub? Stents glabrous, bearing numerous pale

lenticels. Branching not dichotomous. Stipules

otten persistent espccially at flowering codes, ±

orbicular, up to 0.8 x 0.8 cm, narrowing at base,

apex rounded, glabrous on both surfaces. L-eaves

împaripinnate, 3-20 pairs of latéral leaflets, total

length up to 6 cm long; pedole and rachis seg¬

ments 0.3-0.6 cm long, glabrous, channelled

above, narrowly winged, the wings extending

0.5 mm on either side of nndline and incurved;

latéral leaflers narrowly clliprical or elliptical,

(0.45-)0.7-l .9 X 0.2-0.5 cm, of a equal size in

one leaf, the largest often near the midpoint of

the rachis, base sessile, t equal, cuneate, apex

acute; terminal leader narrowly elliptical to

elliptical, 0.7-2. 3 X 0.25-0.8 cm, scarcely larger

than the largest laterals, base sessile or pctiolul-

atc, the petiolule ca. 0.3 cm long, apex acute;

bladc glabrous, coriaceous, cuticle on uppet sur¬

face thick and shiny, not punctate below; margin

recurved, entire or sparsely crenate-denrate with

0-7 notches on each side in the latéral leaflets; on

upper surface, midrib and secondary veins depress-

ed into cuticle and tertiary venation obscure; on

lower surface, midrib somewhat prominent.

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in Sulawesi and Philippines

Fig. 3.—A-D Weinmannia furfuracea A, flowering shoot showing 2 latéral tetrads in sequential leaf axils where in both cases, the

bud between the central racemes Is contlnuing to grow vegetatively; the apical bud ot the shoot is also continuing to grow vegetative-

ly; B, bisexual tlower; C, fruit just before dehiscence; D, seed, (A. B, Tanlra 1595\ C. D, uan Balgooy 3464) — E-J, Weinmannia

eymaeana: E, tlowehng shoot showing an inflorescence of a pair ol dyads, note apical bud between peduncles ot dyads starting to

develop; F, leaf with narrowly winged rachis; G, male flower; H, dehisced capsule; J, immature seed. (E-J, Eyma 3578). —Drawn by

C. Monniè.

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

Inflorescence a pair of opposite dyads; apical

bud glabrous, sometimes continuing to grow

vegetarively during flowering and sometimes

more than one node on a shoot producing dyads

simultaneously; peduncles 0.1-0.3 cm long,

racemes np to 8 cm long, small bud présent in

angle between racemes at apex ol peduncle, glab¬

rous; peduncles t glabrous, axes of racemes

puberulent; bracts at apex ol peduncle persistent,

± triangular. Flowers inserted in fascicles; floral

bracts carinate, sparsely hairy on abaxial surface,

up to 1 mm long, sometimes persistent; flowers

apparently unisexual, only male flowers seen:

pedicel 1 -2.3 mm long, glabrous or puberulent;

calyx lobes 0.7 X 0.5 mm, glabrous; petals

oblong, 1.4-1.6 X 0.6 mm, ± emarginate at apex;

dise lobes 0.3 mm long, narrowly oblong; fila¬

ments up to 2.9 mm long; ovary ca. 0.4 mm

long, pubescent; styles 0.1 mm long, incurved.

Capsules 3.5-4 X 1.8 mm plus style up to

2.2 mm long (just prior to dehiscence), the exo-

carp densely pubescent; calyx lobes and dise

lobes persistent; central column about half the

length of the valves and weakly developed. Seeds

immature, fiat, ca. 0.6 mm long, comose at both

ends.—Fig. 2, 3l>]

Bref.DING SYSTEM. —Polygamodioecious? Male

flowers and fruits présent on Eyma 3578 but not

on the same twig.

FlELD CHARACTERS.—Stems and pedicels red,

flowers pinkish white.

Distribution and ecology. —Endémie to

Sulawesi and known from only 2 collections,

both from Gn. Lumut (Loemoet). Eyma 3607a

was collected on the top of Gn. Loemoet, and

therefore at about 2200 m.

Paratype. — Sulawesi: Eyma 3607a, Res. Menado,

Ond. afd. Poso, Tusschen Biv. 111-Biv. IV, top Gn.

Loemoet, fl., 4 Sep. 1938 (BO).

Eyma 3578 was included by Bf.RNARDI (1964)

in Weinmannia urdaneiensis and appears to be

the basis for his Tab. 29 (except g), but its similar-

ity to W. urdanetensk is only superficial. The

leaves of W, eyrnaeana are in fact more similar to

those of Pancheria multijuga Guillaumin trom

New Caledonia. As sometimes occurs in other

species of Weinmannia, the leaflet blades and

calyx lobes are pustulate.

5. Weinmannia coodei H.C. Hopkins, sp. nov.

A Weinmannia lurfuracea H.C. Hopkins caulibus

juvenibus glabris, non villosis-tomenrosis , et foliolorum

pagine abaxiali glabra non furfuracea recedit.

Type. — Coode 6197 , Sulawesi Tcnggara (SE

Sulawesi), Koaka area, 3“34'S-121 U 40’E, Gn.

Watuwila foorhills above Sanggona, Gn. Sopura,

1600 m, (t., 5 Nov. 1989 (holo-, I.!; iso-, A!, K!).

Small trec 6 m high, 35 cm dbh. Young stems

glabrous, ridged, nodes thickened witb annular

leaf scars prominent, older stems with longitud¬

inal fissures and numerous white lenticels;

branching not dichotomous. Stipules not seen,

caducous. Leaves imparipinnute, with (1)2 pairs

of latéral Jeaflets, total length up to 13 cm; pétio¬

le 1.4-2 cm long, rachis segments 1-1.5 cm long,

pétiole and rachis segments semiterete and flarten-

ed or channellcd and glabrous or tomentose on

adaxial surface; latéral leaflets narrowly elliptical,

distal leaflets larger than proximal ones, the larg-

est per leaf 4.8-6.4 x 1.4-2.1 cm, the base un-

equal, ± sessîle, actlte, apex acute to obtuse;

apical leaflet elliptical to obovate, 5-6.2 X 1.4-

2.3 cm induding base attenuate into a petiolule

ca. 1 cm long, apex obtuse; blades subcoriaceous,

glabrous above and below, not punctate; margin

crenulate, 9-18 notches on each side of the larg-

est latéral leaflets; midrib indented above, pro¬

minent below and glabrous, secondary and

tertïary venation t fiat above and somewhat rais-

ed below, réticulum t dense.

Inflorescence a pairs of dyads; peduncles up to

l mm long, glabrous, the pair of stipules at the

apex of the peduncle sericeous and persistent,

and partially fused at their latéral margins to

form a cup-shaped structure (Fig. 4A’)> each stip¬

ule rounded at apex, abaxial surface strigose,

adaxial surface glabrous; buds at apex of

peduncles in angle between central pair of

racemes not seen; axes of racemes puberu lotis, up

to 7.5 cm long. Floral buds inserted in small fas¬

cicles; floral bracts ca. 0.7 mm long, carinate,

pilose on outer surface. Flowers with pcdicel 1-

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in Sulawesi and Philippines

Fig. 4.—A-D, Weinmannia coodei A, shoot with one ot a pair ot opposite leaves and a developing inflorescence ot 3 racemes; A’,

enlargement of shoot apex (A) showing opposite pétioles and sear from base of interpetiolar stipules, dormant apical bud In centre,

and in axillary positions, between the apical bud and the pétioles, the cupped bracts from within which the racemes arise (as 2 dyads

with one raceme mlssing?); B, flower; C, capsule at dehiscence; D. seed. (A-D, Coode 6197 ).—E-K, Weinmannia hutchinsonii E,

flowering shoot slrowlng an Inflorescence ot 4 dyads, F, stipule; G, male flower; H, female flower; J, capsule at dehiscence, note per¬

sistent dise lobes but calyx lobes fallen; K. seed. (E, F, J. K, Sulit PNH 2781\ G, Elmer 14228\ H, Elmer 17293 ).—Drawn by

C. Monnié.

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

1.5 rnm long, puhetulous; calyx lobes 0.7 X

0.6 mm, pubescent on oucer surface; corolla irre-

gularly obovatc to oblong, 1.4 X 0.8 mm, apex

rounded, margin mïnutely ciliate; dise lobes 0.3-

0.4 mm long, oblong or somecimes fused into an

almosr continuons dise; filaments up to 1.6 mm

long; ovary 0.5-0,7 mm long, densely pubescent;

styles 1,3 mm long, straight; stigmas capitate,

papillose.

Capsule 3.5-4 X 2 mm at dehiscence, the styles

1.5-1.8 mm long, the exocarp densely pubescent;

calyx lobes persistent. Immature seeds 0.6-

0.8 mm long, either comose at both ends or with

hairs ail over surface, Iongest at ends.—Fig. 2,

4A-D.

BREEDING SYSTEM. —Not determined.

FlELD CHARACTERS. —Flowers pink with white

stamens.

Distribution and ecology. —Known only

from the type, front the SE peninsula of

Sulawesi, from mossy ridge forest at 1600 m, on

schist substrate with deep leaf-litter, canopy at 6

m; associâtes include Pandanus.

This species is named after Mark J.E. Coode

who collectée! the type.

6. Weinmannia celebica Koord.

Meded. Lands Piantentuin. 19: 640 [450] (1898);

Koord.-Schum., Syst. Verz. 3: 51 (1914); Engl., Nat.

Pflanzenfam., cd, 2, 18a: 256 (1930); Bernardi, Bot.

Jahrb. Syst. 83. 165 (1964).—Type: Koorders 18022b,

North Sulawesi, Residentie Menado, Loeloeraboelan

nr. Pahoc oere, 700 ni, 8 Apr. 1895 (leao-, here desi-

gnated, BO|; isolecto-, BO!, 12),

Tree 5-27 m, up to 45 cm dbh. Young stems

woody, pilose, the hairs up to 0,5 mm long,

older stents more sparsely pilose to glabrescent

with minute longitudinal fissures and nunterous

lenticels. Stipules few in mature foliage, suborbic-

ular, up to 0.45 cm diameter, base constricted,

abaxial surface strigosc cspeciaily at base, [.caves

imparipinnate with 3-6 pairs of latéral leaflets,

total lcngth up to 14 cm; pétiole 1.5-2 cm long,

rachis segments 0.8-1.1 cm long, petiole and

rachis segments semiterete, somewhat flattened

on adaxial side, pilose and/or puberulent, often

densely so on adaxial side; latéral leaflets lanceo-

Jate or narrowly elliptical, the largest 3-5 X 1-

1.6 cm, the base unequal, + sessile, apex

acuminate; apical leaflet narrowly elliptical, 5.2-7

X 1.4-2 cm, base attenuate into a petiolule ca.

0.7 cm long, apex acuminate; leaflet blades chart-

aceous to subcoriaceous, glabrous above, pilose

below, not punctate; margin crenulate, 11-14

notches on cach side of lhe largest latéral leaflets;

rnidrib slightly depressed aboyé, puberulent at

base, prominent below and pilose, secondary and

terriary venation slightly raised on both surfaces,

réticulum not dense.

Inflorescence a pair of dyads; peduncles laterally

flattened, ca. 1 cm long, strigose-puberulous;

apical bud between the central pair of peduncles

seticeous; bud at apex of pedunde in angle bet¬

ween central pair of racemes sericeous, minute;

axes of racemes pubetulous, up to 11 cm long.

Flowers inserted in fascitles; floral braers ± ligul-

atc, c-a. 1.3 X 1 mm, minutely sttigose, mostly

caducous; flowers probably unisexual; pedicel

1.8-2 mm long, puberulous to glabrous; in old

female flowers/young fruit; calyx lobes 0.8 x

0.6 mm, glabrous; corolla oblong, ca. 1.5 x

0.8 mm, apex rounded; dise lobes 0.2 mm long,

oblong; filaments 1.6 mm long; ovary 1.2 mm

long, densely pubescent; styles 1.3 mm long,

straight or divergent; stigmas papillose. Mature

fruits not seen.—Fig. 2.

JUVENILE FOLIAGE, —Stipules orbicular to

amplexicaul, up to 1.2 x 1.1 cm, persistent.

Leaves imparipinnate, up to 30 cm long, latéral

leaflets in up to 16 pairs, to 5.2 X 1.4 cm, blades

chartaceous, margin distincily serrate.

BREEDING SYSTEM. —Probably dioecious.

PlELD CHARACTERS. —Young leaves red; flowers

dirty white.

Distribution and ecology. —Known from

the northern peninsula of Sulawesi around

Menado in forest at 600-1500 m. Scarce to local-

ly common.

Materiai. EXAMiNF.n.— North Sulawesi: Koorders

18017b, Residentie Menado, Lolomboelan. 900 m, st.,

12 Mar. 1895 (BO. L); Koorders 18021b, Gn. Klabat,

600 m?, st., juv., 19 (an. 1895 (BO); Koorders 18023b,

Oerwoud Lolomboelan bij Pakoe-Oere, 700 m, st.,

juv., 6 Apr. 1895 (BO); Koorders 18025b, Sapoetan-

ADANSONIA, sér 3 • 1998 • 20(1)

Weinmannia in Sulawesi and Philippines

gebergte, 1500 m, st., juv., 5 May 1895 (BO).

LOCAL NAMES.—Teregkoese.

Of the fivc collections cited by KoORDERS-

SCHUMACHI R, only one is fertile anti threc are of

juvénile foliagc. This spccies is distinguishcd

front W fraxinea by ihe pilose indumenrum on

the undersidc of the leaflets and front W elevogel-

ii by thç longer pedicels. 1rs afFmitics seem ro be

with W. hutcbimirnii and W Ittzoniensh front the

Philippines but better collections are needed to

détermine wherher it is in fact worrhy of spécifie

rank.

BernaRDI (1964) cited 2 collections for this

species [Neth hui. For. Serv. bb 20787 (A) front

Sulawesi and Rutten 2231 (U) front Seram] but

did not see the type. Both of these specimens are

now placed in W. furfuracea.

7. Weinmannia negrosensis Elmer

For synonomy and description, see under

Philippines (p. 62).

Materiai. examined.—Norih Sulawesi: Milliken

1034X , Duntoga Boite National Park, Distr. Bolaang-

Mongondow, nr Gn. Sinonibayttga, 0°28’N-

123°44’E, 1970 ni. fl, 25 Sep. 1991 (K).

FlELD NOTES. —Common, multi-stemmed tree

in mossy montane forest; bark white, inner Ltark

orange-brown turning bright orange; wood

white. Petals and anthers white; pedicel, calyx

and pistil pink.

Uncertain and littlc known species

Weinmannia sp. (Sulawesi A)

Monod de Froideville 119, South Sulawesi, afd.

Mandur, Mt, Mamboeliling, N of Mantasa,

2700 m, sr, (BO, L). Alrhough placed by I.AM

(1945) in Weinmannia urrianetensis, its leaves are

quite different, the leaflets being broader, petiol-

ulate, and with rite margin ntore sinuate.

Shrub or small tree; young stems densely

pubescent, branching not dichotomous. Stipules

± orbicular, up ro 0.6 mm wide. apex rounded,

adaxial surface glabrous, abaxial one strigose,

especially towards the base. Leaves imparipinnate

with 9-11 pairs of leaflets, leaves up ro 6 cm

long; petiole and rachis segments terete, densely

pubescent. Leaflets oblong or somewhat ovate,

0.7-1.1 X 0.5-0.6 cm, rhe terminal leaflet not

ntarkedly differing from rhe tarerais in size or

shape, base petiolulate, the petiolule ca. 0.5 mm

long, apex obtuse; blades subcoriaceous, glabrous

above and below except for strigose indumentum

on midrib below; margin crenare-sinuare wirh 3-

4 notches on cach side of a leaflet. Flowers and

fruits not known.

WEINMANNIA IN THE PHILIPPINES

Five species occur in the Philippines, three of

which are endémie. The two which also occur

outside the Philippines are Weinmannia negrosen¬

sis (also in Sulawesi) and W. urdanetensis (also in

New Guinea). Flic ntost wîdcspread Malcsian

species, W. fraxinea, has not been recorded from

this région. Fxcept for a small number of recent

collections, field notes are oftett poor and thus it

is difficult to judge whether the apparent ecolog-

ical différences between species hâve any signific-

ance.

For several of the names published by El.MER

and by MERRILL, the protologucs do not specify

where the holotype is located. According to VAN

StEENIS-KrüSEMAN (1950), the First sets for

these authors should be in Manilla. However,

although the herbariunt at PNH was destroyed

during the Second World War (I lüLMC.REN et al.

1990), some ELMER types for names in

Weinmannia are still extant there (unpubhshed

notes of R.D. HOOGLAND ai P). In these inst¬

ances, the sheets at PNH are provisionally cited

as holotypes. Where a specimen is not known

front PNH, the collections seen or known to

exist are listed as isotypes, and I hâve refrained

from clesignaring lecrotypes because I hâve not

had the opportunity to visit PNH and verify that

no other type materiai is there.

The four species with medium-sized leaflets

( Weinmannia hutchinsonii, W. negrosensis, W.

ADANSONIA. sèr. 3 • 1998 • 20(1)

Hopkins H.C.F.

luzoniensis and U'.' lucida) appear closely related

to one another. Chatacters thar they share indu-

de: calyx lobes frequently Mailing in fruit (which

is unusual in sect. Fasciculata)-, dise lobes short,

broad and persistent in the fruiting stage: flowers

unisexual; capsules densely pubescent; inflores¬

cences often richly developed; stipules often per¬

sistent.

Key to the species in the Philippines

1. Leaves imparipinnate, the latéral leaflets 5-11 pairs, small (0.7-1.5 x 0.25-0.45 cm) .

...........5. W. urdanetensis

1’. Leaves simple, trifoliolate nr imparipinnare, the latéral leaflets up to 7 pairs, the leaves or largest latéral

leaflets per leaf 2-1 5.5 X 0.6-6.5 cm ...... ...2

2. Yoiirtg stems and leaf axes pilose ....... . 3

2’. Ynung stems and leaf axes usunlly glabrous or if indumentum présent, puberulent not pilose .4

3. Litcral leaflets (3-)5-7(-8) pairs, terminal Icaflet 4.1-8.3 X 0.9-2.3 cm; length : breadth ratio for largest

latéral leaflets per leaf I : 0.2-0.3 .1. W. hutchinsonii

3’. Latéral leaflets 2-3L4) pairs, terminal icaflet 6.7-12 x 2,2-5 cm; length : breadth ratio for largest latéral

leaflets per leaf 1 : 0.33-0.52 . 2. W. luzoniensis

4. Leaves simple or trifoliolate, the blades coriaceous, elliptical or obovate, réticulum often dense .

................3. W. negrosensis

4’. Leaves imparipinnate, ( 1 -)2-3 pairs of latéral leaflets, blades subcoriaceous, elliptical, réticulum lax ..

................4. W. ludda

1. Weinmannia hutchinsonii Merr.

Philipp. J. Sci. Bot. 2: 275 (1907); Merr., Enum.

Philipp. Fl. PL 2: 224 (1923); Engl., Nat.

Pflanzenfam., cd. 2, 18a: 256 (1930); Bernardi, Bot.

Jahbr. Syst. 83: 169, t. 20 (1964). -Type: Merrill

5753, Philippines, Mindoro, Mt. Elalcon, Nov. 1906

(iso-, Fl, K!, NY. P!, US)

Weinmarmià tamiguinensis Elntet, Leafl. Philipp. Bot.

7: 2607 (1915); Merr., Enum. Philipp. El. PI. 2:

224 (1923); Engl. Nat. Pflanzenfam., cd. 2, 18a:

256 (1930).—Type: Flmer 14226, Philippines,

Mindanao, Island of Camiguin, Mambajao, densely

wooded guiches or ravines ai 3500 ft. (holo-, PNH,

seen by R.D. HoouiANt); iso-, AI, BMI, BOI, GH!,

K!, L!, P! and numerous others cired by Bernardi).

Weinmannia bidusancnsis Eltner. Leafl. Philipp. Bot.

10: 3723 (1939); nom. illeg., description in

English. Collection cited: Elmer 17293, Philippines,

Luzon, Prov. Sorsogon, lrosin (Mt. Bulusan), 1000

ft., Sep. 1916 (A!. BMI, BO!, GH!, L!, P! and

others).

Weinmannia irnsinensis Elmer, Leafl. Philipp. Bot. 10:

3725 (1939); nom. illeg., description in English.

Collection cited: Elmer 14918 , Philippines, Luzon,

Prov. Sorsogon, lrosin (Mt. Bulusan), 1500 ft.,

Nov. 1915 (BMI, BO!, GH!, L!, P! and others).

Tree 7-12 m high, i 2-40 cm dbh. Young stems

woody, pilose, the hairs up to 0.8 mm long,

older ones more sparsely pilose to glabrescent or

puberulous, with minute longitudinal fissures

and numerous lenticels; branching not dichoto-

mous. Stipules caducous or not, suborbicular or

reniform, up to 1.2 X 1.9 cm, base usually

constiicted, apex roundcd, wavy or coarsely

toothed, adaxial surface glabrous except at base,

abaxial surface puberulous especially at base.

Leaves imparipinnate with (3-)5-7(-8) pairs of

latéral leaflets, total length up to 14 cm includ-

ing a pétiole 1.3-2.5 cm long; rachis segments

0.6-1.5 cm long; petiole and rachis segments

semiterere, flattened on adaxial side, pilose

and/or puberulent, often densely so on adaxial

side; latéral leaflets narrowly elliptical, increasing

in size distally along the rachis, the largest per

leaf 2-6.5 X 0.6-1.4 cm, the base unequal, ± sess-

ile, apex narrowly acute; apical Icaflet narrowly

elliptical. (2.2-)4.1-8.3 X (0.7-)0.9-2.3 cm, base

shortly attetiuate to petiolulate, apex narrowly

acute; Icaflet blades chartaceous to subcoriac-

eous, glabrous above, sparsely pilose below, not

punctate; margin crenate to serrate, 7-11 notches

on each side of the largest latéral leaflets: midrib

slightly depressed above, puberulent at base, pro-

minem below and pilose, secondary and tertiary

venation slightly raised on both surfaces, réticul¬

um not dense.

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in Sulawesi and Philippines

Fig. 5.—Distribution of Weinmannia hutchinsonii.

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

Inflorescence of 1-3 pairs of dyads, sometimes a

few sequenrial nodes producing inflorescences

simultaneously; peduncles larerally flartened,

0.4-1.5 cm long, pubefulous; buds at apex bet-

ween rhe central pair of pedundes I or 3, seric-

eous; buds ar apex of peduncle in angle between

central pair of racemes sericeous, minute; axes of

racemes densely puberulous, up to 12 cm long.

Floral buds inserted in lascicles: floral bracts not

seen, caducous. Flowers unisexual; pedicel 0.8-

1.5 mm long, puberLiJous; calyx lobes 0.5-0.7 X

0.4-0.5 mm, hirsute on ourer surface; corolla

obovate or almost circulai - , 1-1.3 X 0.7-0.9 mm,

rounded at apex, ciliate; dise lobes 0.2-0.3 mm

long, broadly oblong; in male flowers: filaments

2.8-3 ntm long, ovary 0.4 mm long, pubescent,

styles 0.1 mm long, incurved; in temale flowers:

filaments ca. 0.9 mm long, ovary ca. 1 mm long,

densely pubescent. styles ca. 0.9 mm long,

straight, stigmas papillose.

Capsule 2-2,5 x 1.4-2 mm at dehiscence» the

styles up to 1.5 mm long, the exocarp pubescent;

calyx lobes often caducous, dise lobes persistent;

central column présent but shorter than valves.

Seeds ca. 0.6 mm long, comose at both ends, the

hairs to 1.7 mm long.—Fig. 4E-K, 5.

Brehding SYSTEM.— Dioecious.

Field CHARACTF.RS.— Bark grey-brown, verti-

cally cracked and with ± vertical fines of corky

lenticels; outer bark soft and somewhat flaky in

patches, elsewhcre thin and not detaching; inner

bark pinkish straw; wood distinctly reddish or

pinkish. Young leaves reddish. Flowers cream,

light brown, whittsh orange, or pinkish; immat¬

ure fruit pinkish red or violet.

Distribution and ecolouy. —Philippines:

Luzon, Catanduanes, Leyte, Negros and

Mindanao. In fbrest on ridgc-s and slopes, includ-

ing open, disturbed areas front 325-1150 m;

common (Mendoza PNH18409).

Materiai EXAMtNi-n.- Luzon: Edetüo LIS 78206.

Prov. Camarines Sur, Mt. Isarog, Dec. 1928 (K);

Edano PNH 34508. Albay Prov., Ml. Malinao, Fr., 3

Feb. 1956 (A, RM. K, KEP, L); El mer 17293 , Prov.

of Sorsogon, frosin, Mt. Bulusan, fl.. Sep. 1916 (A,

BM, BO, G11, K, L, P); Mendoza 1366. PNB 18409 ,

Albay Prov., Mayon Volcano, 760 m, fr., May-June

1953 (A, K, L); Ramos BS 23494. Prov. of Sorsogon,

yfr., Juiy-Aug. 1915 (A, RISH, BM, BO, GH, K, L,

SING); Ramos BS 23695. Prov. ol Sorsogon, Bulusan

Volcano, fr.. Sep. 1915 (BM, K, P); Su/ir PNH 2717,

Sorsogon Prov., Mt. Bulusan, 380 m. fl., yfr.. July-

Aug. 1947 (A, RO, K, L, SING); Sulit PNH 2781,

ibid., 390 m. yfr., Julv-Aug. 1947 (A. BO, 1., SING);

Vidal 2717, Prov. Canaan tics, Mt. tsarog, fr., Mar.

1886 (A, K).— Catanduanes: Ramos BS 30537, yfr.,

14 Nov-ll Dec. 1917 (A, K), —MindorO; Coode

5393, N coast, Subaan R. inland from San Teodoro,

450 m, yfr., 18 Apr. 1986 (A, K, L).— Negros:

Edano PNH 6811, Oriental, Inalacan R., W of

Tanjay, 500 m, fr.. Sep, 1948 (A, BO, SING),—

Leyte: Wenzel 997, fl., 19 Aug. 1914 (GH); Wenzel

1088, fl., 1 Oct. 1914 (A, BM, GH).— Mindanao:

M.S. Clemens 519, Camp Keithley, Lake Lanao, fl.,

May 1906 (BO, K, P).

LOCAI. NAMES.—Torog-torig (dialect Bic);

Payvagta (dialect Bir).

Weinmannia celebica may be close to this spe-

cies but better collections of the former are need-

ed to détermine whether they are distinct.

2. Weinmannia luzoniensis S. Vidal

Revis. PI. Vase. Filip.: 125 (1886); Merr., Enum.

Philïpp. Fl. PI. 2: 225 (1923); Engl., Nat.

Plktiw.cnlâin., ed. 2, 18a: 256 (1930); Bernardi, Bot.

fahrb. Svst. 83: 172 (1964).—Type: Vidal )i Soler

314, Philippines, Prov. Tayabas, l.ttcban (lecto-. Itéré

deslgnateel, MA, photo at P!; isolecto-, A!, I l, L!, MA

[photos at P!)).

Weinmannia luzoniensis S. Vidal var. puberuht Elrner,

Leafl. Philipp. Bot. 8: 3078 (1919). — Type: Elrner

18066 , Philippines, Prov. Eaguna, Luzon, Mt.

Maquiling, Los Banos, June-July 1917 (holo-,

PNH, seen by R.D. Huotll A,Ntt: iso-, A, BM!, BO!,

K!, 12, P! and others).

Tree 8-20 m high, 10-50 cm dbh. Young

woody stems pilose, the hairs up to 0,8 mm

long, older ones more sparsely pilose to glabresc-

ent or puberulous, with minute longitudinal fiss¬

ures and numerous lenticels; branching not

dichotomoLis. Stipules caducous or not, suborbic-

ttlar, up to 1.7 X 2.2 cm, base constricted, apex

rounded, both surfaces puberulous especially

towards the base. Leaves imparipinnate, wirh 2-

3(-4) pairs of latéral leaflets, total length up to

19 cm including pétiole of 2-2.5 cm; rachis seg¬

ments 0.5-1.5 cm long, petiole and rachis seg-

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

ments semiterete, flatTened on adaxial side,

usually somewhat pubescent ar pilose ac leasr on

the adaxial side; latéral leallets elliptical, increas-

ing in size distally along the rachis, the largest

4.5-8.4 X 1.5-3.5 cm, the hase unequal, + sessile,

apex acuminate; apical leaflet elliptical or usually

markedly rhomboidal, 6.7-12 X 2.2-5 cm, base

shortly attenuatc to form a vvinged petiolule up

to 1 cm long, apex acuminate; leaflet blades

chartaceous to subcoriaceous, glabrous and

sometimes shiny above, sparsely pilose below, not

punctate; margin markedly crenate, 13-17

notches on each side ol the largest latéral leallets;

midrib slightly depressed above, puberulent at

base, prominent and pilose below, secondary and

tertiary venarion slightly raised on both surfaces,

réticulum not dense.

Inflorescence usually 4 dyads or tetrads, some¬

times a fcw successive nodcs producing partial

inflorescences simultaneously; peduncles lateraüy

flattened, puberulous; buds at apex of main stem

between the central pair of peduncles 1 or 3,

sericeous; buds at apex of peduncle in angle bet-

ween central pair of racemes sericeous, minute;

axes of racemes densely puberulous, up to 9 cm

long. Floral buds inserted in fascicles; floral

bracts not seen, caducoils. Flowers unisexual;

pedicel 0.8-1.5 mm long, puberulous; calyx lobes

0.6-0.7 X 0.4-0.6 mm, hirsute on outer surface;

corolla oblong or irregularly obovate, 1-1.2 X

0.6-0.8 mm, rounded or emarginate at apex,

ciliate; dise lobes ca. 0.3 mm long, broadly

oblong; in male flowers: filaments 2.2-3.4 mm

long, ovary 0.4-0.6 mm long, pubescent, styles

0.1-0.4 mm long, incurved; in female flowers;

filaments 0.7-1.2 mm long, ovary 1-1.2 mm

long, densely pubescent, styles 0.9-1.1 mm long,

straight, stigmas capitale, papillose.

Capsule 2-2.7 X 1.3-1.5 mm at dehiscence, the

styles up to 1.5 mm long, the exocarp densely

pubescent to velutinous; calyx lobes caducous or

not, dise lobes persistent; central column présent

but shorter than valves. Seeds ca. 0.7 mm long,

comose at both ends, the hairs to 1 mm long.—

Fig. 6A-C, 7.

Breeding SYSTEM.—Dioecious.

FlELD CHARACTERS. —Flowers yellowish or red

( Conklin & Buwaya PNH 80387).

Distribution and ecoi ogy.— Luzon and one

record from Mindanao. In mid-mountain and

ridge lorest, and secondary r forest, from 500 to

2000 m, on clay soil. Brown (1919) records it on

Mt. Maquiling in mid-mountain forest at 700 m

xs amongst the more prominent first story species

in the Quesxus-Neolitsea association, and says it

also occurs in dipterocarp forest and is fairly com-

mon in some localities in the mountains of Luzon.

Ma i i kia t i xamini.d,— Luzon: Brown BS 18999,

Laguna Prov., Mt. Makiling, fl., Apr.-May 1913 (A,

K) : Conklin (3 Rosarto, PNH 72666, Mountain Prov.»

Mt. Ihukakan, Banane, buds, II., I Aug. 1961 (A, K,

L) ; Conklin dr Buwaya PNH 80387, Mountain Prov.,

Bayninan, Banane, Ifugao, 4000 ft., II., 29 Mar. 1963

(A, K, L); Curran PB 5070, Benguet Pmv., Baguio,

fl.. yfr„ Aug. 1906 (BO, SING); Flmer 8831, ibid.,

fl.. Mat. 1907 (A. BO, K. L); Limer 9093. Prov.

Tayabas, Lucaban, il.. May 1907 (A. BO. K, L);

plmer 18024, Prov. Laguna, Los Banos, Mt.

Maquiling, fi„ June-July 1917 (BM. BO, K, L, P);

Giiatd PB 27274. Benguet Subprov,, Irisait, Fr.. May

1918 (BO, L, SING); Hancock 82, nr, Baguio, st.,

Aug, 1935 (K); Lasasca FB 30194, Benguet Prov., fl.,

Jujy-Oct. 1925 (BO, P); Lan classé 70, Font of Mt.

B.majao, fr., 26 Oct. 1894 (P); Leano FB 21846,

Benguet Subprov., fl., yfr., Jan.-May 1914 (BM, K,

P): l.oher 51/4, Benguet (K); Lober 5125 , ibid. (K);

Merrill Phil. PL 830, Benguet Subprov., fl.. May 1911

(K); Merrill Phil. PL 1739, Benguet Subprov., fl.,

May 1914 (BM, BO. GH, L, P, SING); Meyer FB

3123, Prov Baiaan, La mao R., Mt. Marivclcs, IL,

May 1905 (K, SING); Hamas BS 20533, Prov.

Laguna, San Anrnnin, fr., Feb. 1913 (BM, K, P);

■Hamas BS 23673. l’rov. Sorsogon, Bulusan volcano,

yfr.. Sep. 1915 (K); Sankuhl376, Benguet Subprov.,

sc., Feb. 1916 (A); Salit PNH 6989, Laguna Prov.,

Mt. Makiling, Nat. Park, 500-700 ni, fr., 1 1 Nov.

1946 (A); Vanoverbergh 1253, Boncoc Subprov., fl.,

25 July 1914 (A, l„ P); Vidal y Saler 2002, Benguet

(K), — Mindanao: Ctemens s.n.. Camp Keithley, Lake

Lanao, yfr., Sep.-Oct. 1907 (BO).

LüCAL NAMES.— Tabangawon (dialect Ifugao);

Tabangawon an pabitong, Bani (Tagalog); Itangan

(Igorot); Saiu (Igorot); Sayo (Vidaly Soler 314).

Local uses: good tanbark (Brown 1954).

Typification. —ln the protologue, Vidal y

S01.F.R cites two collections, 314 Lucban, Pr.

Tayabas, and 2002 Distr. Benguet. His original

set of material in Manda was destroyed by fire

and the remainder of his herbarium and types are

ADANSONIA. Sér. 3 • 1998 • 20(1)

Weinmannia in Sulawesi and Philippines

Fig. 7.—Distribution of Weinmannia luzoniensis and W. urdanetensis.

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

at MA (STAFXEU & COWAN 1986). There are

nine sheets of 314 at MA, and the one bearing a

fruiting specitnen and two labels with the in¬

scription “Inspection Gral de Montes de

Filipinas” is here designated as tbe lectorype.

I agréé with BERNARDl (1964: 172) that var.

puberula does not merit récognition.

3. Wcinniannia negrosensis Ëlmcr

Leatl. Philipp. Rot. 2: 577 (1909); Merr., Enunt.

Philipp. Fl. Pl. 2: 225 (1923); Engl., Nat.

Pflanzenfam., ed. 2, 18a: 256 (.1930); Bernardi, Bot.

Jahrb. Sy.sr. 83: 174, l. 23 var. negrosensis (1964).—

Type: Ëlmcr 9656, Philippines, Prov, ofNegros

Oriental, Dumaguetc, Cuernos Mrs,, ridge at 4250

fr., Mar. 1908 (iso-, BM!; alsn A, E, FI, G, LE, and Z

fide Bernarhi).

Weinmatinm simplidfolis Merr., Philipp. j. Sci. Bot.

12: 268 (1917); Merr., Enuin. Philipp. Fl. Pl. 2:

225 (1923); Engl., Nat. Pflanzenfam., ed. 2, 18a:

256 (1930).— WeinrftHrmitt negrvsÇmis var. simplid-

folui (Merr.) Bernardi, Bot. Jahrb. Syst. 83; 175, t.

24 (1964).—Type: Riifnos or F.dano RS 26531,

Philippines, Tayabas Prov., Mr. Dingalan, 300 m, 9

Sep. 1916 (iso-, A, K!. US).

Weinmttnnici cunMtijrilia Engl,, Nat. Pflanzenfam., ed,

2, 18a: 252 (1930).—Type: Ramas Philipp. PL

1287, Philippines, Mindanao (holo-. B, seen by

R.D. HOOGIAND; iso-, BM!, L!, P! and others).

Shruh or tree 3-25 m high, up to 30 cm dbh

when 12 m high. Young stems glabrous, older

ones with numerous white letuicels; branching

not usually dichotomons. Stipules caducous or

not, suborbicular, up to 1.5 X 1.7 cm, base

constricred, apex rounded, abaxial surface glab¬

rous or shortly srrigose especially towards the

base, adaxial surface glabrous. Leaves simple or

trifoliolate; toral length up to 19 cm in trifolîol-

ate leaves, including petiole of 1-4 cm; petiole

subterete, slightly flattened or cbannelled on

adaxial side, glabrous or puberulent; leaf(lei)

blades elltptical to obovate; in trifoliolate leaves,

latéral leaflets 4.5-11.5 X 1.6-3.4 cm, base short-

ly attenuate, apex acute or acuminate; apical leaf-

let 6-15.5 X 2.4-6 cm, base long attenuate

(constricced région up to 1.7 cm long), apex

acute or acuminate; unifoholate leaves 4.5-13 X

1.7-6.5 cm, base attenuate (constricted région

0.6-1.5 cm long); leaflet blades coriaceous, glab¬

rous on both surfaces, not punctatc below; mar-

gin sometimes minute!)' thickened and revolute,

crenate, 11-15 crenations on each side of a leaf¬

let; midrib sometimes slightly depressed above,

prominent below, glabrous, secondary and tert-

iary venation Hat or raised above and raised

below, réticulum dense.

Inflorescence usually 4 dyads or tetrads, some¬

times a few successive nodes producing partial

inflorescences simultaneously; peduncles 0.3-

1.3(-3) cm long, puberulent; buds at apex of

main stem between central pair of peduncles 1 or

3, sericeous; buds at apex of peduncle in angle

between central pair of racemex sericeous, minu¬

te; axes of racemes puberulent, up to 12 cm long.

Floral buds inserted in fascicles; floral hracts

ligulate, 1.5 mm long, ciliate. Flowers unisexual;

pedicel 1-1.7 mm long, puberulent; calyx lobes

0.6-1 X 0.6-0.8 mm, glabrous on outer surface,

ciliate; petals obovate, apex rounded, 1-1.4 X

0.7-0.9 mm, ciliate; dise lobes 0.3-0.4 mm long,

broadly oblong; in male flowers: filaments 2.7-

3.2 mm long, ovary ca. 0.5 mm long, pubescent,

styles 0.1-0.2 mm long, incurved; in female

flowers: filaments 0.9-1.7 mm long, ovary 1.1-

1.4 mm long, pubescent, styles 1.3-1.9 mm long,

straight, stigmas capitale, papillose.

Capsule 2-2.5 X 1.3-1.5 mm at dehiscence, the

styles 1-1.5 mm, the exocatp pubescent or dense-

ly so; calyx lobes usually but not always persist¬

ent, dise lobes persistent; central column présent.

Seeds 0.8-0.9 mm long, comose at both ends,

the hairs to ca. 1 mm long.—-Fig. 6D-K, 8.

BreEDING SYSTEM.—Dioecious.

FlELD CHARACTERS. —Bark greyish or light

yellow-brown; sntooth or finely cracked. Inner

bark red-brown, wood pale straw or “sappy" red,

moderately hard, odourle.ss. Young leaves red-

dish; old leaves bt ight red. Inflorescence axes red-

dish; flowers flagrant; corolla whice tinged with

red, filaments reddish towards the base; or flow¬

ers yellowish white.

Distribution and ecology. —Sulawesi (1

collection) and Philippines (Luzon, Mindoro,

Sibuyan, Ncgros, Leytc and Mindoro) Recorded

from upper montane forest and mossy forest at

1200-1960 m. In the type description, ELMER

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in Sulawesi and Philippii

Fig. 8.—Distribution of Weinmannia negrosensis and W. lucida.

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

records it from dense shrubberies on a wind-

swept ridge at 4250 ft. (= 1400 m).

Materiai. EXAM1NED.—LüZONî Bnrden FB 789,

Prov. Bataan, Lamao R., Mr. Mariveles, buds, May

1904 (BM, K); Borden FB 1227, ibid., fl., June 1904

(BM, K, P); Celestino PNH 7894, Mountain Prov.,

Mt. Polis, Ifugao, 2042 m?, fl., Mar. 1948 (A, L);

Lober 5989, Rizal Prov., Montalban, fr., July 1905

(K); Lober 5990, Rizal, Orind?, fl., 5 Oct. 1906 (K);

Lober 12219 , Rizal Prov., fr., Dec. 1910 (A. BM, P);

Lober 12810, ibid , fl., Mar 1909 (BM); Meyer FB

2756, Prov. Bataan, La mao R, Mt Mariveles, fl., fr.,

Feb. 1904 (K); Quisumbirig <5 Su lu BS 82442,

Benguet Subprov., Mt. Singakalsa. Mar. 1931 (A);

Rarnos BS23500, Prov. Sorsogon, fr., luly-Aug. 1915

(A, K, P); Rames BS 23860, Prov. Sorsogon, Bulusan

Volcano, fl., Sep. 1915 (BM); Rarnos BS 33566,

Ilocos Morte Prov., Mt. P.ilitnlim. fl„ Aug. 1918 (A);

Robinson BS 9399, Prov. Tayabas, Infiinra, fr., Aug.

1909 (L, P); Wbilford 420, Prov. Bataan, famao R.,

Mt. Mariveles, fl., june 1904 (K, P); Wbilford s.n.,

Prov. Bataan, Lamao R., st., May 1905 (K).—

MlNDORO: Coode 5717, NE Mindoro, Ramayan, Mt.

Halcon coinplex above Paitau on Dulangan R.,

1200 m, st.. 8 May 1986 (L).—SlBUYAN: Argent &

Reynoso 89125, Romblon Prov., above Magdiwang on

ridge Icading to Mayos Peak, 1350 m. fl., 27 Aug.

1989 (K).— Negros: Edano PNH 21958, Ncgros

Occidental, Mt. Canlaon, I960 m, st., 10 Apr. 1954

(K, L).— Leyte: RW/ 1057, fr., 14 Aug. 1914 (A,

BM).'— Mindanao: Co 3135, N Cotabato Prov.,

Kidapawan Muncip., Mt. Apo, NW slope, trail bet-

ween Pake Ago and Apo Geothermal projcct site B,

1320 m, buds, 24 Oct. 1990 (A); Ritmos Phil. PL

1287, Camiguin, fr., Apr. 1912 (BM, L, P).

Local names.— Basikong (Bagobo language);

Tangolamos-itum (Visayan).

Typification. —EnüLER did nor cite a particu-

lar collection in rhe protologue of Weinmannin

cuneatifoLia, but only rhe localiry, Mindanao. On

the basis of this, tbe Ramos collection at B is

considered to be the holotype.

Bernard! (1964) recognised two varieties

which had either simple or trifoliolate leaves: var.

simplicifolia and var. negrosensis, However, several

specimens hâve both types of leaves so var. sim¬

plicifolia is not maintained here.

4. Weinmannia lucida Merr.

Philipp. J. Sci. Bot. 10: 7 (1915); Merr., Enum.

Philipp. Fl. PI. 2: 225 (192.3); Engl., Nat.

Pflanzenfàm., ed. 2, 18a- 256 (1930); Bernardi, Bot.

Jahrb. Syst. 83: 170, t 21 (1964).—Type; Ramos

Philipp. PL 1109, Philippines, Luzon, Prov Laguna,

Dabican, in forest along rivet, 17 Sep. 1912 (iso-,

US!; also FI, G, JF). M, U and Z, fide BERNARD!).

Shrub or small tree 3-10 m high. Young stems

± glabrous, older ones glabrous with minute

longitudinal fissures; branching not dichoto-

mous. Stipules caducous or uor, suborbicular, up

to 0.7 X 0.9 cm, base constricted and puberulous

on abaxial surface, glabrous on adaxial side, apex

rounded. Leaves imparipinnate, latéral leaflets

(1-12-3 pairs, rotai Jengrh up to 18 cm including

petiole of 1.5-3.5 cm; rachis segments 1.2-3 cm

long, petiole and rachis segments subterete,

slightly flattened or cbannelled on adaxial side,

sometimes detisely puberuleirt; leaflets elliptical

to broadly elliptical, often conduplicate when

dry; latéral leaflets 5-8.5 X 2-3.7 cm, tbe blade

narrowing uttequally at the base into a petiolule

ca. 0.5 cm long, apex acuminate; apical leaflet 7-

9.5 X 2.5-3.5 cm, not markedly larget than the

largest lacerais, base attenuarc to form a petiolule

ca, I cm long, apex acuminate; leaflet blades sub-

coriaceous, glabrous and sbiny on both sides,

drying dark brown above and reddish brown

below, not punctate; margin sometimes minutely

revolute, crenate, 8-10 notches on each side of a

leaflet; midrib slightly depressed above, prominent

below, secondary and tertiary venarion slightly

raised on both surfaces, réticulum nor dense.

Inflorescence usually 4 dyads or tetrads, some-

times a few successive nodes producing partial

inflorescences simultaneously; peduncles 0.5-

I cm long, .shortly puberulous; buds at apex of

main stem between central pair of peduncles 1 or

3, sericeous; buds at apex of peduncle in angle

between central pair of racemes sericeous, minu¬

te; rachises puberulous, up to 12 cm long. Floral

buds inserted in fascicles; floral bracts caducous.

Female fltnvers (Sulit PNH 6329)'- pedicel 3 mm

long, puberulous; calyx lobes 0.6 X 0.5 mm, glab¬

rous; corolla oblong, 1.1 X 0,8 mm; dise lobes

0,3 mm long, broadly oblong; filaments 1 mm

long; ovary 1.1 mm long, densely pubescent;

styles 1.4 mm long, straight; stigmas capitate,

papillose.

Capsules 2.5 X 1.5 mm at dehiscence, the styles

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in Sulawesi and Philippines

ca. 1 ram, rhe exocarp densely pubescent; calyx

lobes caducous, dise lobes persistent; central

column présent but shorter than valves. Seeds ca.

0.8 mm long, comose at both ends, the hairs to

1.5-2 mm long.—Fig. 8.

BREEDING SYSTEM. —Apparently dioecious.

Distribution and ecology. —Luzon and

?Samar. Edano BS 76049 gives “summit of

forest at 2000 ft. [= 650 m], plant 3 m X 12 cm

dbh, flower whitish pink”. Sulit PNH 6329 (W.

cf. lucida) is from dipterocarp forest at ca. 230 m

on the island ol Samar and is omitted from the

distribution map.

Materjal examiner,—Luzon: Edano BS 76049,

Camarines Sur. Mt. Madooy, 2000 ft., fr., 10 Nov.

1928 (A, SING); Lober5991, Montalban, Rizal, fr., 5

Nov. 1906 (BO, K); Ramas BS 23806 Prov. Laguna,

San Antonio, y fl., Qct. 1915 (A).

W . cf. lucida. — Samar: Sulit PNH 6329, Bagacay,

Concord, 230 m, fl., Apr.-May 1948 (A, 1).

This is a poorly defined species intermédiare

between Weinmannia negrosensis (from which it

differs by having less coriaceous leaves, the réticul¬

um of tertiary and quaternary venation never as

strongly pronounced, and the leaves never

simple) and W luzonicnsîs (which has pilose indu-

mentum on the stems and leaf rachises, the termi¬

nal leaflet are ± rhomboidal, and the margin is

more distinct!}' crenate). However, field observat¬

ions and better ecological information are requir-

ed before W. lucidti can either be equated with

another taxon or more clearly distinguished.

5. Weinmannia urdanetensis Elmer

Leafl. Philipp. Bot. 7; 2608 (1915); Merr., Enum.

Philipp. Fl. PI. 2: 225 (1923); F.ngl., Nat.

Pflanzenfatn., ed. 2, 18a: 256 (1930); L.M. Perry, J.

Arnold Arbor. 30: 160 (1949); Bernard!, Bot. Jahrb.

Syst. 83: 181 (1964) excl. t. 29.—Type: Elmer 13701 ,

Philippines, Mindanao, Prov. of Agusan, Cabadbaran

(Mt. Urdanera), Sep. 1912 (iso-, A. BMI, BO!, BISH!,

K!, IJ and others).

For synonomy, description and illustration, see

New Guinea (Part 3, Hopkins 1998b p. 74).

Material from the Philippines closely resembles

some collections from Irian Jaya.

Distribution and ecology (Fig. 7).—

Philippines (Luzon and Mindanao) and New

Guinea. In Philippines, known from only three

collections. Clemens 16904 describes it as a

“summit rree or high shrub”. In New Guinea, it

grows in montane forest at 1000-3250 m, at

higher altitude in the Central Highfands.

Material examined.— Luzon: Clemens 16904,

Isabela Prov., Mr. Moises, st., Apr. 1926 (SING);

Ramos BS 33268, llocos Norre Prov., Mt. Palimlin,

yfr., Aug. 1918 (A, K, P).

Acknowledgements

This worlt was fïnanced by the Fluman Capital and

Mobility Programme of the European Union via the

network on Botanical Diversity or the Indo-Pacific. I

thank the authoiities of the herbaria cited for access to

spedmens, the late R.D HOOGLAND for his generos-

iry in giving me access to his unpublishcd notes, P.

Lowry and an anonymous reviewer for comments on

the manusetipt, J. FLORENCE for help with the Latin

diagnoses, E. ÀRMÈDE for help with typing and C.

MONNIt: for the illustrations.

REFERENCES

BERNARD! L. 1964.—Revisio generis Weinmanniae.

Pars 111: Sectiones 111 ■ (V - V -VI. Bot. Jahrb. Syst.

83: 126-184; 185-221.

BradIORD J.C. in press.—A cladistic analysis of.spe¬

cies groups in Weinmannia (Cunoniaceae) based on

morpholugy and inflorescence architecture, Ann.

Missouri Bot. Gard.

Brown W.H. 1919.— The Végétation of Philippine

Mountains. Bureau of Science Publication No. 13,

434 pp., Manila.

Brown W.H. 1954.— Usefùl Plants of the Philippines

2: 66. Bureau of Printmg, Manila, Dept. Agtic. &

Nat. Res.

Holmcren P.K., Holmgren N.Ff & Barneït I..C.

(cds.) 1990.— Index Herbariorum, 8 th ed., Part I.

Rcgnum vegetabile, vol. 120.

HOPKINS H.C.F. 1998a.—A révision of Weinmannia

(Cunoniaceae) in Malesia and the Pacific. 1.

Introduction and an account of the species of

Western Malesia, the !,csscr Sunda Islands and the

Moluccas. Adansonia sér. 3, 20: 5-41.

HOPKINS H.C.F. 1998b.—A révision of Weinmannia

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

(Cunoniaceae) in Malesia and the Pacific. 3. New

Guinea, Solomons Islands, Vanuatu and Fiji, with

notes on the species of Samoa, Rarotonga, New

Caledonia and New Zealand. Adansonia , sér. 3, 20:

67-106.

Hopkins H.C.F. & Florence J. 1998.—A révision

of Weinmannia (Cunoniaceae) in Malesia and the

Pacific. 4. The Society, Marquesas and Austral

Islands. Adansonia, sér. 3, 20: 107-130.

Lam H.J. 1945.—Contribution to our knowledge of

the Flora of Celebes (Coll. C. Monod de Froideville)

and some other Malaysian islands. Blumea 5: 554-

599.

Stafleu F.A. & Cowan R.S. 1986.— Taxonomie

Literature, ed. 2, vol. VI: Sti-Vuy. Regnum vegetabi-

le, vol. 11 5.

Steenis-Kruseman M.J. VAN 1950.—Cyclopedia of

collectors. Flora Malesiana ser. l,vol. 1: 1-606.

Manuscript received 30 October 1997;

revised version accepted 27 February 1998.

ADANSONIA, sér. 3 • 1998 • 20(1)

A révision of Weinmannia (Cunoniaceae)

in Malesia and the Pacific. 3. New Guinea,

Solomon Islands, Vanuatu and Fiji, with notes

on the species of Samoa, Rarotonga,

New Caledonia and New Zealand

Helen C.F. HOPKINS

Laboratoire de Phanérogamie, Muséum national d’Histoire naturelle,

16 rue Buffon, 75005 Paris, France.

Masons Arms, Hutton Roof, via Carnforth. Lancashire. LA6 2PE, U.K.

With the collaboration of R.D. HOOGLANDf

Laboratoire de Phanérogamie, Muséum national d'Histoire naturelle,

16 rue Buffon, 75005 Paris, France.

and J.C. BRADFORD,

Missouri Botanical Garden, P.O, Box 299,

St Louis, MO 63166, U.S.A.,

and Washington ünlverslty, St Louis, U.S.A.

KEYWORDS

Weinmannia,

Cunoniaceae,

New Guinea,

Solomon Islands,

Vanuatu,

Fiji,

New Caledonia.

ABSTRACT

Species of Weinmannia bclonging to rwo sections occur in eastern Malesia

and the western Pacific: sect. Fasciculam is largely Maksian, extending as far

east as Fiji, and sect. I.eiospermum is largely Pacific and occurs as far west as

Papua New Guinea. In total, ahout 20 species occur in the région from New

Guinea to the Cook Islands. Weinmannia froxinea, which is widcspread in

Malesia, also occurs in New Guinea and the Solomon Islands. Weinmannia

crofiii from Papua New Guinea (Bismarck Archipelago and Karkar Island) is

described as new, and the new combination W. ouaiememis is published for a

species in New Caledonia. There is a high level of endemism within the

various island groups of this région, although the status of species from

Samoa and the Cook Islands lias not bccn resolved. Régional keys, illustra¬

tions and distribution maps are provided.

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

MOTS CLÉS

Weinmannia,

Cunoniaceae,

Nouvelle-Guinée,

Iles Salomons,

Vanuatu,

Fidji.

Nouvelle-Calédonie.

RÉSUMÉ

Plusieurs espèces de Weinmannia, appartenant à deux sections, existent dans

l’est de la Malésie et l’ouest du Pacifique. La sect. Fasciculata, en grande par¬

tie de Malésie, s’étend vers l’est jusqu’à Fidji ; la sect. Leiospermum, essentiel¬

lement du Pacifique se trouve vers l’ouest jusqu'à la Papouasie-

Nouvelle-Guinée. Au total, environ 20 espèces se trouvent dans la région

comprise entre la Nouvelle-Guinée et les Iles Cook. Weinmannia fraxinea,

largement répandu en Malésie, existe aussi en Nouvelle-Guinée et aux Iles

Salomons. Weinmannia croftii de Papouasie-Nouvclle-Guinée (Archipel

Bismarck et lie Karkat) est une nouvelle espèce décrite ici, et une nouvelle

combinaison, W. ouaiemensis, est proposée pour une espèce de Nouvelle-

Calédonie. Le taux d'endémisme au sein des divers groupes d’îles de cette

région est très élevé, bien que le statut des espèces de Samoa et des lies Cook

n’ait pas encore été résolu. Des clés de détermination régionales, des illustra¬

tions et des cartes de distribution sont présentées.

INTRODUCTION

This is rhe third part of a révision of the

Malesian-Pacifk species of Weinmannia. Part I

(HOPKINS 1998a) gives a general introduction,

including définitions of rhe rerms used for the

structure of the inflorescence, and describes the

species of western Malesia. Parts 2 and 4

(Hopkins 1998b; Hopkins & Florence 1998)

deal with the species of Sulawesi and the

Philippines, and the central Pacific respective])'.

The Weinmannia species dealt with here show

considérable island or island-group endemism,

although 3 number which occur on volcanic

islands and belong to sect. Leiospermum appcar

to be dosely related to one another, and the mot-

phological différences between them are relative-

ly small In addition to the new species and new

combination published here, incomplète material

from Irian Jaya and the Solomon Islands (includ¬

ing Bougainville which is politically part of

Papua New Guinea) appears to represent a hand-

ful of undescribed taxa.

I. WEINMANNIA IN NEW GUINEA AND

THE BISMARCK ARCHIPELAGO

Four named species occur in this région,

Weinmannia fraxinea, W. urdanetensts, W. pullei

(ail sect. Fasvieulata ) and W. croftii (sect.

Leiospermum). Weinmannia pullei and W. croftii

are endémie. Sonic small-leafleted collections

Irom hian Jaya may represent additional taxa but

better collections are needed, and they are not

included in rhe kcy.

Key to the species of New Guinea and the Bismarck Archipelago

1. Flowers inserted on inflorescence axes in fascicles, i.e. a group of pedicels subtended by each bract; inflor¬

escence usually composed of 1-3 pairs of dyads or tetrads inserted in the axils of the most distal pair of

Icaves; apical bud ol shoot, between central pcduncles of the partial inflorescences, présent and usually

dormant ....2

1’. Flowers inserted on inflorescence axes singly, i.e. each individual pedicel subtended by j bract; inflores¬

cence usually a central triad or pentad, developing from the apical bud of the shoot .4. W. croftii

2. l.ateral leaflets in l-4(-6) pairs, 3-10 X 0.8-3 cm; mature seeds comose at each end; inflorescence usually

of 1-3 pairs of dyads or tetrads . 1. W. fraxinea

2’. Latéral leaflets in 1-19 pairs, 0.6-3.2 x 0.3-1.5 cm; mature seeds bearing hairs ail over surface; inflorescen¬

ce usually a pair of dyads .......3

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in New Guinea and the western Pacific

3. Latéral leaflcls oblong or elliptieal, 2-19 pairs; secondary veins oriented at 90° ro main vein; leaflets insert-

ed at 90" to leaf rachis, often strigose-velutinous on underside; calyx pubescent . 3. W. urdanetensis

3’. Latéral leaflets obovate, 1-6 pairs; secondary veins oriented at an acute angle to main vein; leaflets inserted

at an acute angle to the leaf rachis, usually glabrous on underside; calyx glabrous . 2. W. pullei

1. Weinmannia firaxinea (D. Don) Miq.

For synonomy, description, illustration and dis¬

tribution map, see part 1 (Hopkins 1998a; 23).

Breedinü SYSTEM. —Flowers usually bisexual in

New Guinea, rarely unisexual (male: Ledermann

9922 , 9784, Henty & Fureman NGF 42671, ail

front West Sepik; female: Stevens LAE 58123 and

Brass 22460).

FlELD CHARACTERS. —Srnall treelet to entergent,

5-35 m, up to 70 cm dbh, trunk rarely with

buttresses up to 8 m. Bark variable, pale to dark

brown or grêy, smooth, cracked, fissured, scaly or

with pustules; inner bark brown or reddish

brown; sap wood white, straw, pink-straw, some-

times turning purple, hard; heartwood pink or

red; sortietimes with sticky exudate. Buds and

inflorescence axes pink. Flowers whitish, some¬

times tinged with pink, red or green, smell fra-

grant or unpleasant.

Distribution and fcoiocy. —Malesia (except

Sulawesi and Philippines) to Solomon lslands.

See HOPKINS 1998a, Fig. 8, p. 29. In New

Guinea, from 10 m (Vogelkop) to 1450 m and

rarely to 2250 m (Mt. Dayman, Milne Bay and

Mt. Talawe, Hast New Britain). Fôund in a wide

range of forest habitats including lowland rain

forest, hill forest, lower montane and cloud forest

up to the border of subalpine grassland on Mt.

Dayman; frequemly from steep slopes and tidge

tops; in primary forest, disturbed forest and

regrowth and dcscribed as characteristic of older

second growth at 350 m; scattered to dominant.

Juvénile plants common on road cuts and open

areas (Morobe, 100-1 50 m).

At low altitude G 200 m), associâtes include

Linaceae and Anhoptera , At mid élévation (300-

950 m) found in hill forest with Castanopsis ,

Agathis, Freycinetia , and Pandanur, also in hill

forest on limestone karst (600 m, New Ireland),

lower montane, moderately dry forest dominated

by Eucalyptopsis (450 m, Normanby Island), and

lower montane forest dominated by Weinmannia

and Alstotüa spectabilis (950 m, New Ireland). At

higher élévations grows with Podocarpns and

Daaydium.

Weinmannia fraxinea is frequemly recorded

frorn lower élévation in New Guinea and the

Solomons than for instance in Java, Sumatra and

the Malay Peninsula though usually from the

same types of habitat (e.g. road cuts and open

areas, hill forest and lower montane forest).

SeLECTED COLLECTIONS (from a total of 63 xtudied

for this région).—I rian Jaya: Aet & Idja-n 806,

Mcmpcrawaja or. Seroei, buds, 16 Sep. 1939 (A, BO,

K, L, S1NG); Dijk bb 30351 . Ond. afd. Seroei, Eil.

Japen, 800 m, si.. 3 Aug. 1939 (A, BO, L); Kanehira

& Hatusima 12797. Boemi, 40 km tnward of Nabire,

300 m, buds, fr., 11 May 1940 (BO, L, type of W.

h\poglnuca)\ Koster BW8112, Div. Flollandia, Bodem

R., 60 km SE from Sarmi, 70 rn, buds, 10 Feb. 1959

(A, L); Lam 1574, R. Manibctamo, nr. Mt.

Doorman, 1450 m, fl., 9 Oct. 1920 (BO, K); Schram

BW 6009, Vogelkop, Beriat, ± 20 km S of

1 eminaboean, 10 m, st.. 19 Apr. 1958 (L); Sijde BW

4071 , llollandia, Cydoop Mts., Bivouac 1, 500 m,

yfr., 12 Sep. 1956 (A, K, KEP, L); Soengeng

Reksodibtfrdjo 386, SE West Irian, lugembk to Opka,

fr., 8 June 1967 (L); Vmk BW8426, Div. Hollandia,

Sidoarsi Mts., ca. 200 km W of Hollandia, 200 m, st.,

juv., 20 May 1959 (L).— Papua New Guinea:

Western: Henty et al. NGF 31797. nr. Ingembit villa¬

ge, 480 ft„ fr„ 8 June 1967 (BISH. K, L); Rids/Lile &

GaLnre NGF33436, Kiunga, 200 lu. fl., 21 July 1967

(A, K. L). West Sepik: Darhyshïre 238, nr. Miwaute

village, Torricelli Mts., Lumi subdistr., 2600 ft., fl.,

23 Aug. 1961 (A. BISH, CANB, K, L); Darbysbire &

Hoogland8374, nr. Wantipi village on Bliri R., Aivape

subdistr., 800 ft., fl., 3 Aug. 1961 (A, BISH, BM,

CANB, K, L); Henty & Foreman NGF 42671,

Kokomo Creek, trib. of Frieda R., Telefomin subdis-

tt„ 2300 ft.. fl., 28 June 1969 (A, BISH, K. L);

Ledermann 10129, I.nrdberg, 1000 m, fr,, 6 Dec.

1912 (K, type W. alla). East Sepik: Ledermann 8172,

Hunstein Mts., 2-300 m, fr., 9 Aug. 1912 (B, type W.

copU'tttella)■ Madang: PulUn 1040, between Aiome

Patrol Post and Togum village, 1000 ft., fl., 23 Aug.

1958 (A, BM, CANB K, L). Morobe Clemens 1107,

Wareo, 2000 ft., yfr., 28 Dec. 1935 (A, L); Crofi &

Lelean LAE 68552, Natter Bay logging area, 93 km

SE of Lae, 100 m, 30 July 1976 (A, K, L); Crofi et al.

LAE 68601, track from Tigedu to Sambiang, E of

ADANSONIA. sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

Mongi R., 25 km E of Finschhafcn. 800 m, lr.. 17

Sep. 1976 (A, BISH, BM, K- L); Hartley 12829,

above Bakaia, ca. 15 miles SE Cîaraina, 3000 ft., fl.,

26 Jan 1964 (A, L, P); Takeuchi 7135, Markliam vil¬

lage along niargin of l.abu swamp, 100-150 in, fr., 4

Julv 1991 ( A, L); Streimann NGF 26)11, Tiaura,

Kipu, 2600 ft., fi., 7 Jan. 1966 (A, K, L)-, Womerslty

NGF 19044, Oomsis logging area, 1800 fr., buds,

yfr., 29 Nov, 1963 (A, BISH, K, L). Oro Pullen

5931, N side of Sihium Range, S of loma, Bariji-

Managalasa. 3500 fi., si.. 9 Sep. 1964 fl.). Milne Bny.

Brass 22460. N slopes Mt. Dayman, Maneau Range,

2250 m, fr., 24 May 1943 (A, L); Smith NGF 1355,

nr. Mapo, 1 100 ft., fr., Mar. 1945 (A, BISH, K, L);

Stevern I.AF. 58123 , Mi. Duau, above Agaun, 1460 m,

fl., 1 Feb. 1973 (A, K, L); Croft LA F 711)1,

Normanby lsland, NE ot Bwasiaiai, 450 ni, fl., fi., 2

Dec. ] 9^7 (A, RfSH, K, !.), Crofi et al. IAE 68620, S

Fergusson lsland, crack between Ailuluai and

Agamoia, 720 m. fr., 3 Nov. 1976 (A, BISH, K, L);

Brass 27428, Misima lsland, Mt. Sisa, N slopcs,

350 m, fl., fr., 20 Julv 1946 (A, K, L); Gideon L.AE

73267, J.igul .1 lsland, Mt. Riu. 600 m, lr., 1 1 Mar.

1979 (BISH, K, 1). New Britain: Frodin NGF

26807 , Mt. Talawe, Talasea, 6400 fi., fr., 25 May

1966 (A. BISH, BM, CANB); Fies étal. NGF34400,

6 miles F of Fullerborn Harbour, 300 in, fr,. 8 May

1973 (A, BISH. K, I ). Sayen NGF 21989, T'irilongi

village, Kandrian subdisti . 1300 fi., fr„ 14 Mar. 1965

(L). New Jreland. Crofi LAt 65576, 5 km S of

Logagon village, N Schleinirz Range, Logagon sub-

distr., 600 m, fr., 23 Oct. 1974 (A, BISH, BM, L);

Gideon LAE 77167, Lelet farm, Lelet Plateau, Konos

subprov., 950 m, fr., 29 Oct. 1984 (A, K, L).

Local nam LS. —Mesjeforon (Tehid language,

Vogelkop); Saboo (Tor language) and Hassip

(Manikiong language) both trom Sidoarsi Mis.;

Tima (Orne language, Wantipi) and Yeb-peh

(Wapi language, Miwaute) both W Sepik;

Yibit(s) (Western Prov.); Suweti (middle Watia,

Morobe); Gabisamina (Upper Waria, Milne

Bay); Vani'ldaidalava (Fcrgusson lsland).

Variation and riïlationshii'S. —A complex

and variable ochlospecies, and several names

from New Guinea hâve been put into synonomy

for the First time (see HOPKINS 1998a, p. 23).

Relatively variable within New Guinea, showing

almost the complète range of variation on this

one island. Thcrc is no morpbological overlap

with Wcinmannia pullei and W. urdanetensis even

though the largest ieaflets ol W. pullei may

approach the size of ïhe smallest orics in W.

fraxinea.

2. Weinmannia pullei Schltr.

Bol. Jalirb. Svst. 52: 164 (1914); Nova Guinea 12:

492, t. 192 (1917); Engl., Nat. Pflanzenfam., ed. 2,

18a; 255 (1930); Bernardi, Bot. Jahrb. Syst. 83: 176,

r. 25 (1964); P. Roycn, Alpine Fl. New Guinea 4:

2539, r. 739 (1983).—Type: Bulle 470, West New

Guinea (Inan Jaya), on surnmit of Mi. Peramcles, ca.

1100 m, 27 Nov. 1912 Choit»-, B; iso-, BMI, BOI, K!,

L!).

Weinmannia virgulata Schltr., Bot. Jahrb. Syst. 52:

164 (1914); Nova Guinea 12: 492 (1917); Engl.,

Nat, Pllanzenfam. ed. 2, 18a; 255 (1930).—Type:

Bulle 692, West New Guinea (Irian Jaya), on Mt.

I leliwig, ca. 1700 tn, 15 Dec 1912 (liolo-, B; iso-,

RO!. L!),

Weinmannia versreegbii L.M. Perry, J. Arnold. Arbor.

30: 162 (1949); Bernardi, Bor. jahrb. Syst. 83: 183

(1964).—Type: Brass & Vemeegh 10469 (Irian

Jaya), 9 km north-east of Lake Habbema, 2700 m,

Oct. 1938 (holo-. A; iso-, BM!, BOI, K!, L!).

Shrub or rree (1.75-)4-27 tn high, up to 30 cm

dbh, rarely epiphytic. Young stems, buds and

young leaves sericeous, older stems glabrcscent,

bearing numerous pale lenticels. Branching not

usually dichotomous. Stipules usually caducous,

ligulate to ± orbicular or reniform, up to 0.8 X

1 cm, narrowing at base, apex roundcd, abaxial

surface strigosc espccially rowards rhe base,

adaxial surface glabrous. Leaves imparipinnate

with l-6(-IO) pairs of latéral Ieaflets, total length

up to 10.5 cm long, induding pétiole 0.7-1 cm

long; rachis segments ca. 0.8 cm long; petiole

and rachis diverging boni the stem at an acute

acroscopic angle, terete or somewhat flattened on

upper surlace jusr below' point ol insertion of

Ieaflets, sometimes winged, rbe wings extending

to 0.8 mm on cither side of mid line, petiole and

rachis, glabrous, or pubcrulous above, or tontent-

ose, the hairs up to 0.5 mm long; latéral Ieaflets

obovate or oblanceolare, 0.6-3. 2 X 0.3-1. 5 cm,

inserted at an acute acroscopic angle to the leaf

rachis, base equal, apex acute to obtuse; terminal

leaflet narrowly elliptical to clliptical, 0.8-4.8 x

0.3-1.7 cm, base attenuate, apex acute; blade fiat

above, usually glabrous on both surfaces, some¬

times sparsely strigose below especially on

midtib, subcoriaccous, not punctate; margin

crenulate or rarely dentale with 3-7 notchcs on

eacb side in the latéral Icafiet»; on upper surface,

midrib and secondary veins ± fiat, secondary

ADANSONIA. sér. 3 • 1998 • 20(1)

Weinmannia in New Guinea and the western Pacific

Fig. 1.—A-H Weinmannia urdanetensis A, tlowering shoot witb Inflorescence consisting of a dyad (base only) and a teirad. B, C,

leaves showing range in leaflet shape and number; D, male flower; E. detail ot D with perianin removed lo show pubescent gynoe-

cium with short styles, minute dise lobes and a single stamen; F temale flower with one petal removed to show short filaments; G,

capsule atdehiscence with persistent calyv and one petal rémaînlng; H. seed. (A, Bowers 401; B, Versteagh BW10312; C, Rabbins

2922, D, E, Hoogland 3 Pullen 5463; F, fiVacte AMU 7663, G, H. Paijmana 131S) —J-Q, Weinmannia pullei J, flowering shool with

an inflorescence consisting of four racemes subtended by a pair ot reduoed leaves, the apical bud between the dyads continuing to

grow vegetatively. and anDther pair of racemes borne on a latéral shoot, also subtended by reduced leaves; K, L. leaves showing

range In leatlel size and shape, M, male llower, N, bisexual flower; P, capsule at dehiscence with persistent calyx, Q, seed. (J,

Hoogland & Schodde 6979 ; K, M, Vink 17098 ; L, l /inas & Wiakabu LAE 59405; N, Womersley NGF 24540 ; P, Q, Hoogland &

Schodde 7685). — Drawn by C. Monnié.

ADANSONIA, sêr. 3 • 1998 • 20(1)

Hopkins H.C.F.

veins at an acure acroscopic angle to the main

vein; midrib prominent below.

Inflorescence usually of 2 dyads, the apical bud

of main stem somerimes aborting or continuing

to grow vegetatively during flowering, or rarely

of 2 or 4 individual racemes; sometimes a few

successive nodes producing dyads simultaneous-

ly; a pair of reduced leaves and a pair of stipules

usually présent at base of the racemes in each

dyad; peduncles 0.6-2 cm long; racemes up to

12 cm long; buds at apex of peduncles between

bases of racemes minute, velutinous, sometimes

starting to grow during reproduction; peduncles

and rachises usually sparsely puberulent. Floral

buds inserted in fascicles, floral bracts elliptical,

offen hairy, 1-1.2 x 0.5 mm, persistent; flowers

mostly unisexual, sometimes bisexual; pedicel

1 .5-3.5 mm long, puberulent or ± glahrous;

calyx lobes ca, 0.5 x 0.4-1.4 x 1.1 mm, glahrous

or with ciliate margin; petals elliptic, 1.1-2.3 X

0.7-1.5 mm, margin minutely ciliolate; dise

lobes 0.3-0.6 mm long; in male flowers: fila¬

ments up to 4.1 mm long, ovary 0.5-1 mm,

pubescenr, styles 0,2 mm long, incurved; in

female flowers; filaments up to ca. 0.8 mm long,

ovary 0.5-1 mm long, pubescent, styles ca.

0.9 mm long, straight, puberulent at least at

base, stigmas capitale, papillose; in bisexual

flowers: filaments ca. 4 mm long, ovary ca.

1 mm long, styles ca. 1 mm long, straight,

Capsules 3-4.5 x 2.5-2.8 mm at dehiscence, the

styles 0.7-1.2 mm long, the exocarp densely

pubescent; calyx lobes persistent; central column

about half length of valves. Seeds ca. 1 mm long,

bearing hairs ail over surface, longest at ends.—

Fig. 1J-Q, 2.

Rri-EI ItNt ; SYSTEM. —Polygamodioecious? Fruits

develop from flowers with both long and short

filaments.

FlELD CHARACTERS. —Outerbark brown, dark

grey or créant, rather smooth with irregular

130° 135° 140° 145° 150°

Fig. 2.—Distribution of Weinmannia pullei.

ADANSONIA, sér. 3 ■ 1998 • 20(1)

Weinmannia in New Guinea and the western Pacific

cracks and lenticels; inner bark brownish, oran-

ge-brown or reddish brown. Wood cream to

pink, of medium hardness, bomogeneous.

Flower buds Jighr grecn; Flowers creamy white,

fragrani: sepals yellowish-green or red; corolla

and filaments white; anthers pinkish-yellow;

ovary light green turning pink, style pink at base

and white at tip.

Distribution and ecology.— Endémie to

New Guinea. Grows as a tree in montane forest

with Nothofagus , Podncarpus, Phyltacladus and

Pandanus and as a small tree or shrub in mount¬

ain top shrubbery; from 1100-2100 m in Irian

Jaya and 2800-3200 m in Papua New Guinea

(PNG). Common,

SelecTF.D COLLECTIONS (front a total of 28 stud-

ied).— Irian Jaya: Eyrna 5207 , Wissel l.akcs région,

Moetaro, 11., 9 Sep. 1939 (BO, L); Eytm 5224, Ibid.,

Tarapadimi, S of Pake Tage Mneraro, buds, 1 Sep,

1939 (L, StNG); Pullc 188, Mt. Perameles, 1100 ni,

fl., 28 Nov. 1912 (K, L).— Papua New Guinea-.

West Senik: Vinas & Wiakabu IAF. 59405 , Tcle-

fomin subdistr., Silinmogu, ridge below Tamanagabip

on track to Busilmin, S°0Q'S-14r 3 05 f, 2800 m, st.,

1975 (A, B1SH, K, L). Southern Highlunds Frodin

NGF26964. N slope Mt. Ne, 9200 h., 26 July 1966

(A, BlSH, K, L); Frodin NGF 28131, nr. Lei cantp on

tack to Mi. Ambria, Tari, 8600 ft., fr., 29 July 1966

(A, L); Gil/hun NGF 25132, Ihiwara, Tari Gap, ,3400

ft„ 11 June 1966 (A, BLSH, K, I.),- Kalkman 4800,

Mt. Ne, Tari subdistr., 3140 m. 12 July 1966 (A,

BlSH, L); Vint 17098 , Tari subdistr., N slope of Mt.

Kerewa, 3015 m, 7 July 1966 (A, BlSH, L, P). Engiv.

Hoogland & Schtrdde 69/9. nr. Poio village, SE ridge

of Yaki valley, Wabag subdistr., 9000 fr., 8 Julv 1960

(A, BlSH. BO, K, L); Hoogland & Schodde 7685,

Yobobos grassland area, source ol Lagaip R., Laiagam

subdistr., 8500 ft., fr., 5 Sep. I960 (A, RM, CANB,

L); Rabbins 3056, S slopes Ambum-Marimuni I livide

nr. Londau village, Wabag area, 8000 ft.. fl„ fr., 24

July 1960 (CANB, L); WomersUy NGF 24540,

Kandep-Lagaip Di vide. Laiagam subdistr., 9600 ft., 1

May 1965 (A, RJSH. CANB, K). Western

Highlunds. Pullen 116, F. rim of M(. Ogu, 12 miles F.

ofMt. Hagen station, 8500 ft., 12 July 1957 (A, RM,

K, L); Robhim 1138, Min) subdistr., Pinj R. valley nr.

Banz R.C. sawmill, 8000 ft., st., ,30 July 1957

(CANB); Veidkamp & Stevem 5493, Klangun Hill, S

of Tomba. 2700 in, 11, 29 Mav 1972 (BlSH, L);

Pullen 51 ? 3, WHP/Simbu, Min)-Nona Divide, N

side of Kubor Range S of Minj, 9540 ft., fl., fr., 20

Aug. 1963 (A, CANB, K, L, P).

LOCAL NAÏVES. —Enga language: Kain, Kain-

raggan-taggan (Poio), Tagantagan and Tagar

(Kepilam); Autiggli, Tewara no. 2 and Mabi

(near Tari); Tandan (Mendi), Kwirap (Minj),

Gubidigiii (Margarima).

Variation. —Material of Weinmannia from

monrane New Guinea with small leaflets can be

divided into two main groups, W. pullei (includ-

ing W. venteeghii with a narrowly winged rachis)

and W urdanetensis. Weinmannia pullei has

rather few, obovare leaflets and W. urdanetensis

usually has more numerous elliptical or oblong

ones and rheir leaflets also differ in their angle of

insertion. These two species appear to be ciosely

related and share the following characters: inflor¬

escence primarily of dyads; floral bracts often

persisting to fruiting stage; Flowers usually uni-

sexual but sometimes bisexuah fruits with the

valves densely pubescent; seeds with hairs ail over

their surface and not conflned only to the ends.

Their distributions arc largcly sympatric but

there is only a small minority of intermediate

collections. The structure of the inflorescence is

partïcularly variable in W pullei.

Mosr collections of W. ptdlei are from Papua

New Guinea and are relatively uniform in leaflet

size, shape and vestiture, although Vinas &

Wiakabu NGF 59405 (Snow Mts„ West Sepik

Prov., Fig. IL) has unusually small leaflets.

Collections from Irian Jaya are more variable:

a) The three collections from near Lorentz River

(Mrs. Perameles and Hellwig) {Pulie 488, 4 7 0,

092) hâve 3-9 pairs of leaflets per ieaf and the

leaflets margins are dentate not crenate in two of

them.

b) In the Wissel Lakes région, Eyrna. 5207 is

vegetatively quite close to many collections from

PNG but Eyma 5224 has trifoliolate Ieaves with

small leaflets that are distinctly hairy on rhe

Jower surface, especially on the main vein.

c) In the Vogelkop, van Royen & Sleumer 7224

and Gjellerup 1159 both hâve largcly trifoliolate

Ieaves (the nuntber ol leaflets varies from 1-4). In

the former, the leaflets are rather rounded at the

apex and not very distinctly crenate; both collect¬

ions are from small shrubs at 2080-2100 m.

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

Weinmannia versteegbii was distinguished by

both Pkrry (1949) and Bl.RNARDl (1964) by its

winged rachis and petiole. Ffowever, rhere is a

continuum in thc shape of the rachis and petiole

within W. ptdlei.

3. Weinmannia urdanetensis Elmer

Leafl. Philipp. Bot. 7: 2608 (1915); Merr., Enum.

Philipp. 1 1. PI. 2: 225 (1923); Engl., Nat.

Pflanzenfam., ed. 2, 18a: 256 (1930); L.M. Petly> I.

Arnold Arlior. 30: 160 (1949); Bcrnardi, Bor, Jalirb.

Syst. 83: 181 (1964) exri. i. 29.—Type: Elincr IJ70I,

Philippines, Mindanao, Prov. of Agusali, Cabadbaran

(Mt. Urdaneta), Sep. 1912 (ko-. A, RMI, BO!, B1SH!,

K!, L! and others).

Weinmannia tricbophora L.M. Pcrry, J. Arnold Arbor.

30: 161 (1949); Bernardi, Bot. jihrb. Syst. 83: 181

(1964).—Type: M.S. Cterne ns 9498 , NE New

Guinca (Papua New Guinea), Morobe Distr., nr.

Samanzing, 2100-2400 m, in mountain bush, 18

Jan. 19,39 (holo-, Al; iso-, B).

Weinmannia nrwoguineensis L.M. Perry, J. Arnold

Arbor. 30: 161 (1949).—Type: M.S. Ctemens 7517,

NE New Guinea (Papua New Guinea), Morobe

Distr., Sarawaket. 1800-2400 m. in mountain

foresr, 9 Nov. 1937 (holo-, A!; iso-. B).

Shrub or tree, 3-26 m high, up to 43 cm dbh,

variable in lorm front bushy to gnarled to slend-

er. Young stents velutinous, axillary buds and

young leaves sericeous, older stems glabrescent,

bearing nunterous pale lenticels. Branching not

usually dichotomous. Stipules usually caducous,

± orbicuiar or reniforfit, up to 0.8 x 1.1 cm,

narrowing at base, apex rotinded, abaxial surface

sparsely to densely srrigose especially towards the

base, adaxial surface glabrous or puberulenr.

Leaves imparipirinate with 2-19 pairs of latéral

leaflets, total lettgth up to 10.5 cm long, inciud-

ing petiole ca. 0.5 cm long; rachis segments 0.3-

1 cm long; petiole and rachis diverging from the

stem at an angle of almost 90" especially at grow-

ing tips; petiole and rachis segments terete, den¬

sely velutinous, the hairs erect, up to 0.5 mm

long; latéral leaflets elliptical, oblong or some-

what avare, the margins ± parallei, 0.6-2.6 x 0.3-

1.1 cm, inserted ± at 90° to leaf rachis, base

equal or unequal, cuneate to square to cordate,

apex broadly acute; terminal leaflet narrowly

elliptical to elliptical, 0.9-3 X 0.3-1.2 cm, base

petiolulate, the petiolule ca. 0.3 cm long, apex

acute; leaflets imbricate or not, fiat or revolute,

usually glabrous or puberulent on upper surface,

the cuticle thick and shiny, sparsely to densely

strigose-velutinous belovv. the midrib velutinous;

blades coriaceous, not punctate; margin crenul-

ate wirh 4-7 notches on each side in the latéral

leaflets; on upper surface, midrib and secondary

veins depressed into cuticle, secondary veins ± at

90° to main vein, tertiary venation obscure,

midrib prominent below.

Inflorescence a pair of dyads (rarely a pair of

tetrads or 4 dyads or 4 individual racemes),

sometimes a few successive nodes producing

dyads simulraneously; the apical bud of the main

stem between the bases ol the dyads densely

vclutinous and often contiuuing to grow vegeta-

tively during flowering; bud at apex of pedimcles

between bases ol racemes minute, velutinous;

peduncles Ü.3-0.4 cm long; racemes up to 9 cm

long; peduncles and axes of rhe racemes densely

velutinous. Flowers inserted in fascicles; floral

bracts elliptical, often hairy, 1.1-2 mm long, per¬

sistent; floWei'S unisexual or rarely bisexual; pedi-

ce! 0.5-1 mm long, puberulent or ± glabrous;

calyx lobes 0.7-1 X 0.6-0.9 mm, hirsute; petals

elliptical, oblong or almost circulai', 1-1.9 X 0.7-

1.2 mm, rounded or notched at apex, margin

minutely çiliolate; dise lobes 0.3-0.6 mm long;

in male flowers: filaments 2.2-3.6 mm long,

ovary ca. 0.6-1 mm, pubescent, styles 0.2-

0.5 mm, incurved; in lettude flowers: filaments

0,8-1.7 mm long, ovary 1-1.5 mm long, pubesc-

enr, styles 0.5-1 mm long, straighr, puberulent at

least ai base, stigmas capitute, papillose; in

bisexual flowers: filaments 2.6+ mm long, ovary

1-1.5 mm long, sryles l-l 8 mm long, straight.

Capsules 2.5-4 x 1.5-2 mm at dehiscence, the

styles ca. 1 mm long, thc exoearp densely

pubescent; calyx lobes persistent; central column

abour half lengrh of valves. Seeds ca. 1 mm long,

bearing hairs ail over surface, longcst at ends.—

Fig. 1A-H, 3.

BREEDING SYSTEM. —Largeiy dioecious but

sorae exceptions. Wa.de ANU 7663 has both male

and female flowers on the same sheet (though

not the same stem) and Womersley NGF 15240

has male, female and bisexual flowers. Robbi?is

ADANSONIA. sér. 3 • 1998 • 20(1)

Weinmannia in New Guinea and the western Pacific

191 has male and bisexuaJ flowers in the same

inflorescence.

FlELD CHARACTERS.—Outer bark grey-brown,

silver-grey or rarely black; inner bark pinkish

brown with fine red streaks or dark srraw brown;

sap wood whire ro pink, pale brown or orange,

very hard; heart wood pink or dark red. Floral

bads green, pink or red; flowers white or cream,

rarely light green.

Distribution and ecology.— Philippines and

New Guinea. In lower inontane and moncane

forest; also secondary fbrest and open scrub on

limestone; ofcen recorded from ridges or on steep

slopes; abundant and gregarious. Associates

include Litsea, Panda nus, Cinnamomum and

Nothofagus. In the Central Highlands of Papua

New Guinea, recorded from 2500-3250 ni bur at

lower élévations elsewhere (e.g. 1000-1900 m in

Irianjaya, 1200 m in Torricelli Mrs., 1500 m in

Hunstein Mrs and 1770 m on Mi. Simpson,

Milne Bay Prov.).

S H F.CTED coi ( GCnoNS (from a total of 47 studied

for tltis région).— Irian Java: Kostenmms 2304,

Vogelkop, Arfak Mrs., Angi gira lake, Manolcwari

subdistr., 1900 m, fl., fr., 9-22 Oct. 1948 (BQ, L);

Versteegh BW 10312, Kcbar valley, Watjetoni Mt.,

1200 m, 23 Nov. I960 (L); Sijde RW 5388, ibid.,

'I’obi Mtb„ 1000 m, 11., 18 Jutic 1958 (L); Brass &

Verstetgb 11908, 15 km SW ol Bernhard Camp,

Idenburg R., 1750 m, fr., I I Jan. 1939 (A, BM, RO,

K, L). Papua New Guinea: West Sepik-, Frodin

UPNCi 42.33, Aitape subdistr., ridge on Aitape side of

Mt. Somero, Torricelli Range, 1200 m, st., 2 Feb.

1974 (K. 1.). East Sepik: Hooglvid & Craveu 10952,

F. ridge of Sumset (Mt. Hunstein), Ambunti subdistr.,

4500 ft., sr., 12 Aug. 1966 (CANB, L); Takeuchi

6340, Hunstein Range, Mt. Sainsai, 1000+ m, fr., 24

July 1990 (A). Madame: Pullen 6131, S slopes

Finistcrre Range to S of Mt. Abilala, 8100 ft., fr., 17

Nov. 1964 (BM, CANB). Southern Highlands : Crofi

et al. LAt 61048, 2 miles N laro R., S slope Mt.

Giluwe, Mcndi subdistr., 2340 m, fr., 21 Jan. 1974

(A. K, L). Enga : Elenley AN U 2391, 4 miles NW of

Kupalis, nr. Wabae. 8400 ft.. buds, 16 Jan. 1965 (A,

K, I.); Hoogland & Scbodde 7254, nr, Kepilan village,

Lagaip valley, Laiagam subdistr., 8000 ft., fl., 30 July

Fig. 3. —Distribution of Weinmannia urdanetensis in New Guinea (see also Hopkins 1998b, Fig. 7).

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

I960 (A, BISH, BM, L); San rider s 1008, R.C.

Mission Timber lease, Wabag-Komiam rd., Wabag

subdistr,, 7900 ft, st., juv., 15 July I960 (CANB, L);

Womersley NGI■' 15240 , Wabag-Laiagam rd., 24

miles, margin L. lviva, 1 mile SE Sirunke, 9000 ft.,

fl., 22 July 1962 (A, K, L). Western Highlands:

Bowers 401 . Hagcn subdistr., Pokaripuku, Kopaka,

upper Kaugel, 7600 ft., fl., fr., 24 Nov. 1968 (I,); van

Royen NGF 18168, confluence of Warapuri 8c Koti

rivers, Wahgi-Jimmi dividc, N of Nondugl, Minj sub-

distr., 7200 ft., buds, 4 Sep. 1963 (A, BISH, K, L);

Saunders 627, lower slopcs of Mi. Hagen. 2 miles

NW of Tomba, 8000 ft., fl., fr., 28 June 1957 (A, L);

Saunders 675, top Mr, Oga, 8500 ft., st, 12 July 1957

(A, L); Vinas l/PNG 4988 , Mi. Ambrangaba, NE of

Milep village, Minj distr. 2350 m. fl., 21 Eeb. 1981

(L); Womersley NGF 5352, Al R., Nondugl, st., 7 July

1953 (A, BO, K, L, SING). Srmbu: Brass 30306, Mt.

Wilhelm, E slopes, 2770 m, fr., 3 July 1959 (A, K, I.);

Sterly 80-40, Chimbu valley, Gurugaro Ongo,

2500 m. fl., 14 July 1980 (A, L); Wade ANU 7663,

Keglsugl area, Upper Chimbu valley, 9000 ft., fl., 20

June 1967 (A, CANB, K, L). Eastem Highlands:

Gmbh & Edwards 391 , W of Farîma R., Marafunga

sawmill, Goroka subdistr.. 2600 m, yfr., 25 July 1971

(L); Hoogland & Pullen 5463, nr. Daulo camp, As.iro-

Mairi dividc, Goroka subdistr., 2500 m, fl.. 26 lune

1956 (A, BISH. RM. CANB, K, L); Marti,y 13274,

Marafunga. 20 miles NW of Goroka, 9500 ft., st., 13

Oct. 1964 (A, CANB, K). Morobe: Hoogland 9668,

Mannasat, Cromwell Mt».. Huon Peninsula, 7900 ft.,

st., juv., 22 Aug. 1964 (CANB); Schadde & Craven

4976, E slope of Spreader Divide, S miles NW of

Aseki, 7400 ft., st., 18 Apr. 1966 (L), Milne Bay\

Pullen 7857 , Mt. Wadimana ridge, NE from Mt.

Simpson, 1770 m, st., 22 Julv 1969 (L).

LOCAL NAMF.S. —Asro (Kebar language,

Vogelkup); Air», Jine, Kain (Wahgi language,

Minj); Dekiso (Naho language, Finisterre Mes.);

Kain-tachai (Enga); Kumare (Chimbu) Kumare

yaundo kembre (Kutnan language, Chimbu);

Kumai (Hagen, Togoba); Dzaliamchgehtalalawa

(Dunantina), Duascgeh (Chimbu, Masul);

Tagantagan (Enga language, Kepilam); Pone-

Kuni (Western Highlands?); Q-anak (nr. Wabag,

Merimanta); Yambabengo (Chimbu); Uspa

(Waskuku, Mt. Hunstein)

Geographicai. VARIATION.— Spécimens from

the western part of the distribution tend to hâve

more numerous pairs of longer, narrower leaflets

(Philippines and Irian Jayà; also Torricellis and

Mt. Hunstein but the latter are stérile). In the

Central Highlands of PNG, leaflets on fertile

specimens tend to be shorter and broader, often

more densely pubescent (“W. trichophora") and

there are fewer pairs per leaf. A number of speci¬

mens from the Central Highlands hâve comp-

aratively large leaflets, approaching W. pullei ;

most of tbese collections are stérile and some

show a range in variation in leaflet size on one

sheet. Fertile collections with large leaflets include;

Robbins 191 , Hoogland & Schodde 7254 and

Womersley NGF 11280.

RelATIONSHIPS,—A pparent ly closely related to

W. pullei from New Guinea and W. clemensiae

from Mt. Klnabalu in Bornéo, which it

resembles in its branching pattern and somewhat

bullate leaflets.

4. Weinmannia croftii H.C. Hopkins, sp. nov.

Weinmannia denhamii Seem, et W. vitiensis Seem.

affinis sed ah ilia foliolis lateralibus uno usque tribus

(non uno usque duadecimis) paribus dispositis, maximis

per folio 2.9-6.7 X 0.9-1 .7 cm (non 1.1 x 3(-4.8) x

0.3-1- J(-1,5) cm) dijfert, et ab bac quae folia trifoliola-

ta foliolis latioribus et magnis cortaceis possédée.

Type.— C.E. Ridsdale NGF 36706, Papua New

Guinea, Madang Prov., Karkar Gland, 4°40’S-

145°57'E. 3000 ft, fl., yfr., 22 Jan. 1968 (holo-, L!;

iso-, AI, BfSH!, K!).

Shrub or tree, 2-20 m high. Young twigs short-

ly hairy, finely ridged with narrow longitudinal

fissures and round or elliprical lendcels; branch-

ing somerimes dichotomous. Stipules usually

caducous on reproductive branches, sometimes

persistent on végétative ones, usually elliptical,

ligulate or rhontboidal, ca. 0.7 X 0.4 cm. obtuse

to broadly acute at apex, + glabrous on adaxial

surface, strigose on abaxial surface, densely so

towards rhe base. Leaves trifoliolate or impari-

pinnale with 1-3 pairs of leaflets, up to 14 cm

long including pétiole of 1.3-2.5 cm; rachis seg¬

ments 0.8-1.7 cm long; pétiole and rachis seg¬

ments semiterete, flattencd above with a narrow

central ridge and winged, rhe wings extending

ca. 1 mm on eitber side, in eath rachis segment

the wings broader distally towards point of insert¬

ion of opposite leaflets; petiole and rachis usually

bearded above, glabrous or bearing a iew hairs on

underside; latéral leaflets lanceolate to narrowly

elliptical, the proximal ones often shorter than

ADANSONIA, sér. 3 ■ (998 • 20(1)

Weinmannia in New Guinea and the western Pacific

the more distal ones when in several pairs, 2.9-

6.7 X 0.9-1.7 cm, apex acute, base asymmetrical,

the distal side acute to attcnuate, proximal side

obtuse; terminal leaflet narrowly elliptical to

narrowly obovate (2.8-)4.3-l 1 X (0.9-) 1.2-

2.3 cm, apex acute, base attettuate; blade glab-

rous on both surfaces, usually punctate below,

subcoriareous to coriaceous; margio somewhat

thickened and minutely revolute, crenate especial-

ly towards the apex with 14-17 crenaüons on each

side; midrib narrowly prominent above and slight-

ly prominent below, sometimes hirsute.

Inflorescence a central triad or usually pentad,

the lower racemes in the axils of leaves or not;

peduncles and rachis segments 0.9-1.6 cm long;

racemes 3-9 cm long; inflorescences axes puberul-

ent. Floral buds inserted singly; floral bracts lan-

ceolate or somewhat carinace, up to 2 mm long,

somewhat strigose, caducous. Flowers unisexual;

pedicel 1-2 mm long, puberulent; calyx lobes tri-

angular, 0.7-1.1 X 0.6-0.9 mrn, ± glabrous; pelais

elliptical, 1.1-1.7 X 0.7-1 mm; dise lobes 0.4-

0.8 mm long; in male flowers; Filaments 2.4-

3 mm long, ovary 0,6-1 mm long + glabrous,

styles minute, 0.2 mm long and curved inwards;

in female flowers; Filaments 0.9-2 mm long,

ovary 1-1.8 mm long, almost glabrous, styles

0.9-1.5 mm long, stigmas capitale and papillose.

Capsules distincdy supported by réceptacle,

valves 2-4 X 1.3-1.8 mm ai dehiscence, the styles

0.8-1.3 mm long; exocarp minutely ridged,

almost glabrous (few minute strigose hairs); calyx

lobes not persistent; central column présent and

often persistent on réceptacle after valves hâve

fallen. Seeds 0 8-1 mm long, 16 per capsule,

comose ar both ends, the hairs ca. 0 5 mm

long.—Fig. 4A-E, 5.

BREEDINC SYSTEM. —Dioecious.

FlELD CHARACTERS. —Shrub in open areas such

as scoria slopes, or a tree in forest. Flowers white

or cream: calyx pale green, corolla and filaments

Fig. 4.— Weinmannia croftii: A, stem showing infructescence (pentad) with reduced leaf at node; B, stipule; C, male flower; D,

female flower; E, dehisced capsule. (A, Foreman & Katik LAE 59132', B, Frodin NGF 26810-, C, Ridsdale NGF 36706', D, Croit &

Katik NGF 14957, E, Vandenberg & Mann NGF 42302). —Drawn by C. Monnié.

ADANSONIA, sér. a • 1998 • 20(1)

KAR KAR

NEW IRELAND

NEW BR1TAIN

£ W. crottn

▲ W. purpurea

| W. ysabelensis

W. sp. (Solomons A)

^ W. sp. (Solomons B)

H* W. exigua

NEW GUINEA

^ca.

BOUGAINVILLE

VELLA LAVELLA

KOLOMBANGARA -

NEW GEORGIA

VANGUNU

SANTA ISABEL

GUADALCANAL

SMALL 3

MALAITA

SAN CRISTOBAL

Fig. 5.—Distribution of Weinmannia in the Bismarck Archipelago and Solomon Islands (except W. fraxinea).

Weinmannia in New Guinea and the western Pacific

white, anthers yellovv oi straw, dise vellow-oran-

ge, ovary liglu green, stigmas white to brown.

Distribution anp ecqlogy.—P apua New

Guinea: Karkar Island, Manus and New Britain.

In New Britain, collected from 1200-2100 m,

where it is an early coloniser on volcanic sub¬

strates as they become stable after an éruption,

and sometimes very abundânt e.g. on Mt.

Ulawon (D. FroDIN pets, comrn.). Also found

on non-volcanic substrâte in mossy montane

Nothofdgus forest where its associâtes înclude

Nastus and Gleichenia, soils indude clay over

Ümestone. On Manus, collected from 530-

720 m and on Karkar, from 820-1050 m in

lower montane ridge forest with Eugenia,

Panda nus-, Dilknia , and Elaeocarpus , and a colon-

ist on lava flows where it was co-dominant with

Dodonaea and Eurya. Not yet recorded from

New Iteland, where Weinmannia fraxinea is

found. Weinmannia crofïii and W. fraxinea are

sympatric only in New Britain.

Parai vi'i s.— Papua New Guinea: Bismarck

Archipelago: East New Britain: Clunie LAE 63017.

nr. latte, central Nakanai plateau, Pomio subdîstr.,

5°32’S-15]“18’E, 1200 m, il.. 28 Nov. 1974 (A,

BISH, K, L); Cruft crKatik NGF 11057. Mr. Suie, ca.

25 miles NNE of Fulletborn Harbour. Pomio subdis-

tr„ 5' > 50’S-15Ü‘5O’E, 1500 m, fr., 8 May 1973 (A, K,

L); Isles et ai NGE34406. Mt. Ltiluiua, ca. 30 miles

NE of Fulletborn Harbour, Pomio subdistr.. 5"45’S-

150°50 > F„ 2000 m, fl., 11 May 1973 (A, B1SH.K, L).

West New Britain: Fradin NGE 26810. Mt. f'alawt,

summit. l alasea, 3 V 32’S-I48“18 L, 0400 h., fl., 25

May 1966 (A, L); StevensLAE 5 J252, NNE slope Mt.

Ulawon, Hoskins subdistr., 5°2’S-15)“22’E, 3600 fr.,

buds, 20 Feb. 1971 (A, K, L); Vinas LAE 59724, Pake

in Nakanai Plateau, F. Nakanai. 5°29'S-151 “ 16’E,

1610 m. 11., fr., 11 Nov. 1975 (A, BISH, BM, K, L).

Manus: Foreman & Knfik LAE 59132, Mt. Dremsel,

Lorengau subprovr., 2 0 I5'S-149°50’E, 600 m, fl., 25

Oct. 1974 (A, L); Kenaga et ai. LAE 77543 , ibici,

2°09’S-146‘ > 56 , 30 I ’E, 630 m, fl., 26 Mar. 1981 (A, K,

L); Stone & Streimann l.A /: 53681, ibici., 2°10'S-

146°55'F, 1800-2000 Et., yfr.. 21 Junc 1971 (A.

BISH, L); Sittttù et al. 2903, Mt. Dremsel, 2”08’S-

146°57'L, ca. 6 km iuland from Pelekawa, fr., 29

Nov. 1975 (K). NE New Guinea: Madang , Karkar

Island. Ridsdale NGE33981. 4°40’S-145°57’E, 2500

ft., fr., 16 ]an, 1968 (A, BISH, K, L): Vandenberg dr

Mann NGF 42302, 4°40’S-146°00’F, 3200 ft., 12

]une 1969 (I,),

Local NAMES.— Naligugu (Frodin NGF26810,

New Britain), Sirikat ( Ridsdale NGF 36706,

Karkar),

Rhl ATIONSHIPS. —Appears to be closely related

to Weinmannia denhamii from Vanuatu and W.

vitiensis from Fiji and possibly conspccific with

Weinmannia sp , A from the Solomons. Ail hâve

imparipinnate, coriaceous leaves with a winged

petiole and predominantly unisexual flowers.

Weinmannia denhamii usually has more numer-

otis, shorter leaflets than W. croftii, and W.

inticnsis has trifoliolate leaves with rather broader

leaflets than those of W. croftii. The three are

kept separate for the présent since the popula¬

tions are disjunct and the mean values of the

foliage characters are different for each species,

even though the ranges overlap.

This species is named after Jim CROFT, former-

ly at LAE,

Insufficiently known species

Weinmannia sp. (New Guinea A)

Lam 1869 (Irian Jaya, Doormantop, 3100 m,

fl,, 30 Oct 1920 [BO, I !]) is from a shrub 0.5 m

high: leaves trifoliolate; leaflets orbicular, round-

çd at apex, coriaceous; margins minutely notch-

ed; leaflets, pétioles and stems dcnscly hairy.

This collection appears to he a Weinmannia but

it does not match any described taxon, and fert¬

ile material is needed hefore it can he named. It

is most similar to van Royen & Steamer 7224

from the Vogelkop (which is placed here in W.

pullei) but that specimen is almost glabrous and

its leaflets are more obovate.

II. WEINMANNIA IN THE SOLOMON

ISLANDS (INCLUDING BOUGAINVILLE)

Six species occur in this région: Weinmannia

pttrpurea , W■ sp. /I and W. sp. B in scct.

Leiospermum-, and W. fraxinea, W. ysabelensis and

W. exigua in sect. Fasciculata. Except for W.

fraxinea, most species are known from fcw collec¬

tions. Bougainville in particular appears to be

ADANSONIA. sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

undercollected and W. fraxinea has not yet been occurs in Fiji (see p. 94 for description) whereas

reported from there. Weinmannia exigua also the other four species are endemic.

Key to the species of the Solomon Islands

1. Flowers inserred sitigly in the Axils of bracts on the axes of the racemes; inflorescence usually a triad or

pentad: if leaves compound, then rachis winged, the wings extending > 0.7 mm on either sidc of midline

of pétiole and rachis ......2

1’. Flowers inserted in small fascicles in the axils of bracts on axes of the racemes; inflorescence of 1 or 2 pairs

of dyads or tetrads inserted in sériés at most distal leaf-bearîng node; leaves compound and if rachis wing¬

ed, then wings extending < 0.4 mm on either side of midlme of petit le and rachis .4

2. Leaves usually simple(rarely trifoliolate) and relatively large ([3-] 4.2-18 X 1.3-5 cm) .... 4. W. purpurea

2’. Leaves trifoliolate or imparipinnate, leaflets < 5.5 X 2 cm.3

3. Latéral leaflets 1-2 pairs, 2.5-3.S x 1 1.5 cm ..........5. W. sp. A

3’. Latéral leaflets 3-5 pairs, 0.7-L3 X 0.4-0.5 cm.<.6. W. sp. B

4. Latéral leaflets (l-)2-4 pairs, 3.5-9 X 1-2.7 cm; rachis rerete to semiterete, not winged . 1. W. fraxinea

4’. Latéral leaflets 1-5 pairs, 1.3-2 X 0.7-0.8 cm; rachis minutely winged (wings extending < 0.2 mm on

either sidc of midline) .. 2. W. ysabelensis

4”. Latéral leaflets 6-9 pairs, 0.9-1.3 X 0.4-0.5 cm, rachis narrowly winged (wings extending < 0.4 mm on

either side of midline) .3. W. exigua

1. Weinmannia fraxinea (D. Don) Miq.

For synonomy, description, illustration and dis¬

tribution map see Hopkins (1998a, p. 23).

BreeDINC. SYSTEM.—Flowers bisexual in the

Solomons, though the anthers are unusually

small (ca. 0.25 mm diameter).

FlELD CHARACTERS,—-Tree up to 40 m high,

often with plank-like buttresses up to 2 m high.

Bark variable: reddish brown and flaking, Jight

brown and fissured, dark brown and scaly, or

smooth. Slasb also variable: slash wood reported

as reddish brown, light brown, pink, white,

whitisb brown or rèd, tbe texture hard, soft, or

coarsely 11 bru us; a detailed description is given

on Walker & White BSIP 150 (A). Buds reddish;

flowers whire (rarely créant, yellow, lighr brown),

scented or not.

Distribution and ecoi.ogy. —Malesia (except

Sulawesi and Philippines) to Solomon Islands.

See HOPKINS 1998a, Fig. 8, p. 29. Not yet recor-

ded from Bougainville or Makira. From 10-

620 m, in well drained primary and secondary

forest, often on ridge tops; also in forest over

coral limestone, including disturbed forest, fiat

plain, and forest-grasshmd boundary.

Material examined.— Solomon Islands:

Choiseul: Gafiii BSIP 18977, SE, W of Oaka River,

1800 ft., fr., 12 Mar. 1970 (L). Vella LavelLi: Kotali

BSIP 11173, S, Oula R. area, 360 ft.. fl., fr., 13 Aug.

68 (K, L). Kolomhangara-, Mauriasi et aï, BSIP8603,

NE. Kokove Area, 1300 ft., fl., yft, 12 Jan. 1968 (K,

LU Mauriasi et ni. BSIP 8687 , N. Shouider Bill Area,

400 li„ ir., 19 Jan. 1968 (K, L); Mauriasi et ni, BSIP

11361. SE, 2000 h., buds, 6 (une 68 (K, L); Mauriasi

et al. BSIP 11493, SE, Shouider Hill Area, 800 fr., fl.,

1 7 lune 68 (K. L); Mattriasi et al. BSIP 11647, NW,

Rei Area, 900 fi., fr., 3 July 1968 (K, L); Whitmore &

Womtrsley BSIP 838, SE, Bambari Harbour, fl., 24

Nov. 1962 (K, L); Whitmore BSIP 1436, W coast,

Merusu Cove, 340 ft., fl.. 14 Feb. 63 (K. L). New

Georgia: Burn-Murdoch 's eollecton BSIP 6939, SE,

Maki, fl., 26 Oct. 1966 (R. L): Cowrnentfow BSIP

3690 , NW, Vaimbu R„ 230 ft. fl., fr., 18 Mar. 1964

(K, 1,): Whitmore's çollccturs BSIP 3737 , NW, nr.

Hovoro, 565 ft., buds, 21 May 1964 (K, L).

Vangutiu: Maenu'u BSIP 6420, Davala, 160 fr., fl.,

22 July 1965 (R, E); Whitmore BSIP 871 , SE coast

between Vura village and Gcvala R., fl., 29 Nov. 1962

(K. L): Whitmore BSIP 946, SE coast between Vura

village and Gevaia R., inland nr. Merusu lslet, 3 Dec.

1962 (R, L), Santa Jsabel: Becr s cuUci tors BSIP 5124,

NW, Kolokofu R„ 20 ft., fr., I Apr. 1966 (K, L);

Susui BSIP 8319, Allardyce Harbour, 30 ft., buds, 17

Nov, 1967 (K, L). Guatlahanal Gafui et al. BSIP

9442, L. ridge S ofMakina Station, 120 ft., buds, fr.,

14 Mar 1968 (K, U; Am- BSIP 4909, NW, Mt.

Mambulu, 1500-1600 ft.. fi., 17 Nov. 1964 (R. L);

Ltpaqeto BSIP 3376, NE, Rere R., ca. 3 miles inland,

fr., 21 Nov. 1963 (K. L); Mauriasi et al. BSIP 11823,

E, Makina Area. 180 ft., buds, yff., 29 Sep. 1968 (R,

L); Mauriasi et ai BS/P 12221 , SW, Wandcrcr Bay

Area, 800 ft., fl., 18 Oct. 1968 (K, L); Nakisis &

Mauriasi BSIP 8122 , NW, E of Mt. Mambulu, 1000

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in New Guinea and the western Pacific

Fig. 6. —A-F. Weinmannia purputea A left sîde ot a flowerlng shoot with opposite umfoliolate leaves and one ot two partial inflor¬

escences, (heptad) that nas reduced leaves at one node: B. stipule: C. lemale flower: D, detail of C with perianth removed to show

gynoecium, dise lobes and one stamen: E, capsule at deniscence with central column; F, seed. (A. C. D, F, Kajeivski 1738 B, E,

Mauriasi BSIP 12092). — G-L Weinmannia ysapelensis G, tlowering shoot with inflorescence of four dyads; note apical bud bet-

ween inner pair of dyads; H. detail of a node showing dormant apical bud and two axlllary buds starflng to develop, and the bases of

two opposite pétioles; note stipular scar beneath apical bud; J, bisexual flower; K, detail of J with perianth removed to show gynoe¬

cium, dise lobes and single stamen; L, fruit. (A-L, Brass 3215). —Drawn by C. Monnié.

ADANSONIA. sér. 3 • 199S • 20(1)

Hopkins H.C.F.

ft., fl., yfr., 14 Nov. ï 967 (A, K, L); Sirute'e et al.

BSIP 9900, SE, nr. Logu Village (Avu Avu), 1900 ft.,

fl., Fr., 1 June 1968 (K, L.); SinUee et al. BSIP 10074.

W coast, Alidavata ( l'alise) Kiki Ridge, Duidui, 1000

ft., fl., 13 June 1968 (K, L); Whitnwre BSIP 1283.

Honiara, Mi. Au,sien, yfr., 31 Dec. 1962 (K, L).

Malaita: Fa’arndo et al. BSIP 13492. NE, 450 ft., fl.,

fr„ 22 Nov. 1968 (K, L); Whiniwre BSIP 3874, SW,

Are Are district, W coast, Kilt, fl., 13 Dec. 1963 (K,

L). Stnall Malaita: Gafui et al. BSIP 16322,

Waiusuusu Area, 170 ft., fl., 26 Aug. 1969 (K, L).

LOCAI. NAMCS. —Aitoto ( Kwara’ae) commonly

recorded; other names are Futankwai and

Wangalu (hoth Kwara’ae names), Oliollmwani.

DISCUSSION. —Although this is the most com-

monly collected species from the Solomons

Islands, most of the collections date from the

1960s and this species was not recorded from

here by Bernardi (1964). Material from the

Solomons is rather uniform.

2. Weinmannia ysabelensis L.M. Perry

J. Arnold Arbor. 30: 162 (1949); Bernardi, Bot.

Jahrb. Sysr. 83, 183 (1964).—Type: Brass 3215,

Solomon Islands, Ysabel, Tiratona, 600 m, mountain

forest, Nov. 1932 (holo-, Al; iso-, BISH, BM!, BRI,

BO!, G, L!. S1NG!).

Tree 25 m tall. Young twigs densely pubescent,

the hairs to 0.4 mm long; older twigs sparsely

pubescent; branching not dichotomous. Stipules

caducous, ca. 3.5 mm long, sericeous, apex

rounded. Leaves imparipinnatc, with 1-5 pairs of

latéral leaflets, up ro 9 cm long, including petiole;

petiole and rachis segments 0.5-1.3 cm long,

semiterete, sparsely hairy along midtine on

adaxial side, hirsute on abaxi'al side, minurely

winged, the wings extending 0.2 mm on either

side; latéral leaflets ovate to oblong, 1.3-2 X 0.7-

0.8 cm, the most proximal ones smaller than

more distal ones, apex broadly acute, base

somewhat unequal to dimidiatus; apical leaflets

narrowly ovatc-elliptical, 3-4.4 X 1-1.3 cm, larger

than latéral leaflets, apex acute, base often

abruptly constricted into decurrent petiolule;

blades glabrous on botlr si des except for hairs ca.

0.5 mm long on midrib on adaxial surface, not

punctate; margin somewhat thickened and cren-

ate, 4-6 or 8-9 fcrenations on each side of latéral

and apical leaflets respectively; midrib minutely

indented above and prominent below.

Inflorescence of 2 or 4 pairs of dyads; apical

bud of the main stem berween the bases of the

central peduncles sericeous; bud at apex of

peduncles between bases of racemes minute,

velutinous; peduncles 0.3-0.7 cm long, racemes

up to 8 cm long; inflorescence axes puberulent.

Flowers ittserted in fascicles or singly, especially

distally. Flowers bisexual pedicel 1.5-2.3 mm

long, sparsely puberulent; calyx ca. 0.5 X

0.4 mm, glabrous; petals obovate or oblong, 1.3

X 0.8 mm. apex rounded or emarginate; dise

lobes up to ca. 0.3 mm long; filaments ca. 3.5

(-4.5) mm long; ovary ca. 0.9 mm long, pubesc¬

ent; styles up to 1-7 mm long; stigmas papillose.

Immature capsules with ovary 2.8-3-5 X

1.5 mm, exoearp pubescent, the styles 0.8-2 mm

long; calyx and dise lobes mostly persistent.

Immature seeds ca. 20 per capsule, comose at

both ends.—Fig. 5, 6G-L.

BrEEDING SYSTEM. —Flermaphroditic.

FlELD CHARACTERS.—Large tree with a thick

bole. Bark reddish brown and flaky, the wood

hard and red. Flowers white, immature fruits

pink.

Distribution and ecology. —Known only

from the type from Ysabel Island where it is des-

cribed as common.

This species appears closely related to

Weinmannia fraxinea although the leaflets are

much smaller, Further collections are needed to

détermine the précisé relationship between these

two species.

PERRY recorded that rhe seeds had hairs ail over

their surface (as in W. pullei and W. urdanetensis)

and not only at rhe ends The holotype from A

has a packet attached with a mature fruit and a

seed that marches Pl-RRYs description. I lowever,

the fruit is much older than those attached to the

twigs with leaves on the sheet, and the seeds the

latter contains are immature and hâve hairs only

at the ends. It seems douhtful therefore that the

seed described by PERRY was aetually lrom this

specimen.

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in New Guinea and the western Pacific

3. Weinmannia exigua A.C. Sm.

For typification, description, illustration and

breeding System see under Fiji (p. 94).

FlELD NOTES. —Common tree up to 20 m high

(Bradford pers. coram.), with dense rounded

crown and llowers in off-white panicles

( Whitrnore BS/P 995)■

Distribution and ecology. —Originally des-

cribed from Fiji where il is known (rom two coll¬

ections. Said to be common on crater rim on

Van(g)unu Island at 2000 ft., on ridge with deep

humus {Whitrnore BS! P 995)- —Fig. 5, 9A-C.

MATERÎAt. I X AM IN CD,—SOLOMON ISLANDS: New

Georgia Group, Whitrnore BS/P 995, Van(g)unu

Island, inland from Merusu Islet, on narrow parts of

ridge with deep humus laver, crater rim, ait. 2000 ft.,

fl., 10 Dec. 1962 (I,!, LAE).

LOCAL NAME. —Wangulu (Kwara'ae) ( Whitrnore

B SI P 995).

Whitrnore BS/P 995 was previously equated

with Weinmannia urdanetensis (Whitmore

1966) which occurs in New Guinea and the

Philippines, but the leaflct shape is not a dose

match, lt is similar to the matcrial from Fiji in

the size, shape and texture of the leaflecs and die

narrowly winged rachis, and agréés well with

Howard 89 in the number of pairs of leaders.

The leaflets are also similar in shape to those of

W. ysabelensis although they are smaller and more

numerous (ca. 8 pairs).

4. Weinmannia purpurea L.M. Perry

J. Arnold. Arbor. 30: 159 (1949); Bernardi, Bot.

Jahrb. Syst. 83: 198, t. 34 (1964). —Type: Kajeunki

1738 (Papua New Guinea). Bougainville Island,

Kupeî Gold Field, 1000 m. tain forest, Apr. 1930

(holo-, A: iso-, BISH. BMI, BRI, G, GH, L!, LE, S,

SING!, US).

Tree 20 m high. Twigs terete, scarcely dattened

at nodes, branching frequently dichotomous.

Internodes up to 8 cm long, Stipules caducous,

elliptical, ca. 0.8 X 0,4 mm, apex rounded or

obtuse, sparsely strigose on abaxial surface, gla-

brous on adaxial one. Leaves usually simple or

unifoliolate, the petiole articulated at jonction

with blade or not, or rarely trifoliolate; simple

leaves broadly lanceolate or narrowly ovate (3-)

4,2-18 x 1.3-5 cm. apex acure to caudate, base

attenuate to decurrent; in unifoliolate and trifo-

liolate leaves, articulated petiole ca. 1.5 cm long,

narrowly winged, sparsely puberulenr; in trifolio¬

late leaves, latéral leallets ca. 3.5 X 1.2 cm, une-

qual at base: leaf blade glabrous on both surfaces,

punctate below, subcoriaccous; margin somewhat

thickcned and minutely revolute, strongly crena-

te ( 13-) 18-25 crenations on each side of the

blade; midrib narrowly cidged above and strongly

prominent below, secondary and rertiary vena-

tion slightly raise on bodt surfaces.

Inllorescence usually a central penrad (some-

times a triad or heptad) ot composed of 2 latéral

partial indorescences. the apical bud aborted;

triads sometimes developing in subdistal nodes

also; peduncle and rachis segments 1.4-4.5 cm

long, racemes up to 17 cm long; inflorescence

axes puberulent to almost glabrous. Floral buds

inserted singly; dorai bracts * linear, 1.8 mm

long, caducous. Flowers unisexual, only temale

llowers scen, the stantens shortly exserted from

rite flower and shorter than the styles; pedicel 1-

2.1 mm long, puberulent; calyx lobes triangular,

0.8 X 0.5 mm, glabrous; pelais ovate, 1.4 X

0.8 mm; dise lobes up to 0.7 mm long; filaments

ca. 1.5 mm long, anther 0.3 X 0.3 mm, entpty;

ovary ca. 1.5 long, sparsely and minutely strigose

or + glabrous; styles 0.9-1.3 mm long; stigmas

capitate and papillose.

Capsules 3.1-3-5 X 1.2-1.5 mm at dehiscence,

the styles 1.1-1.3 mm; exocarp with minute

longitudinal ridges, almost glabrous with a few

minute strigose hairs, warty; calyx lobes not per¬

sistent; central column présent. Seedsca. 0.8 mm

long, 16 per capsule, comose at both ends, the

hairs ca. 1 mm long.—Fig. 5, 6A-F.

Breeding System. —Dioecious?

Field cuaracters,— Bole crooked, ca. 50 cm

dbh, but tresses absent. Bark lighr brown smooth;

slash: wood hard, reddish brown, bark soft.

Flowers very pale purple on purple stems

(Kajewski /738) or white with a faint smell

(BSIP 12092). Fruit purple (Kajewski 1738),

ADANSONIA. sër 3 • 1998 • 20(1)

Hopkins H.C.F.

green when immature (BSIP 12092).

Distribution and f.cology. —Known from

only t\vo collections, growing at 785-1000 m.

Habitats described as rain forest and well drained

primary (orest on ridge top. Common according

to Kajhwski.

Materiai examinf.d.—Papua New Guinea:

Bougainville : Kajeivski 1738 (BO, L, P, SING,

type).— SOLOMON ISLANDS: SW' Guadalcanai .

Mauriasi BSIP 12092, Duidui area, 2400 ft., fl., yfr.,

9 Oct. 1968 (K, L).

Local NAME. —Aitoto (Kwara’ae).

Uncertain and little known species

5. Weinmannia sp. (Solomons A)

Weinmannia sp,, Corner, Philos. Trans. Ser. B 255:

577 (1969).

Tree up to ca. 8 m high. Stems glabrous,

branching sometimes dichotomous. Stipules

caducous, iigulate, ca. 4 X 2 mm, with short

strigose hairs on the abaxial surface. Leaves tri-

foliolate or imparipinnate with 2 pairs of latéral

leaflets, up to 7.5 cm long, including the pétiole;

pétiole and rachis segments 1-2 cm long, glab¬

rous, winged, the wings extending up to 1 mm

on cither side of midline; latéral leaflets elliptical,

2.5-3. 8 X 1-1.5 cm, base ± unequal, apex acute;

terminal leaflets elliptical, 4.5-5.5 X 1.7-2 cm,

base attenuate, apex acute; blades glabrous on

both surfaces, punctate below, coriaceous; mar-

gin crenate (but much chewed by insects).

Inflorescence composed ot triads or pentads,

sometimes a few successive nodes bearing partial

inflorescences simultaneously, and reduced leaves

at nodes within the inflorescence. Floral buds

inserted singly; floral bracts ca 2 7 mm long,

exceeding buds, Iigulate, minurely strigose.

Pedicel puberulent; calyx lobes glabrous. Mature

flowers and fruits unknown.—Fig. 5.

Immature foliacé. —In Corner RSS 1188, the

leaves hâve up to 3 pairs of narrowly elliptical

leaflets; young stems, leaf axes and underside of

the midrib are puberulent. Hill RSS 9011 has

broadly ovate latéral leaflets and the axes and

leaves are glabrous.

FlELD CHARACTERS.—Bark and wood red.

Inflorescence in bud red; flowers pinkish ( Corner

RSS 89) or whitc and swcetly fragrant (J.C.

Bradforo pets. comm,).

Distribution and ecology. —Collected at

700-900 m on Vangunu (J.C. BRADPORD pers.

comm,), 1100 1800 m on Kolombangara

(BRADFORO pers. comm. and Corner RSS 89)

and 2340 m on Guadalcanai (Hill RSS 9011).

Habitats include forest on summit plateau and

ridge of mountains, Common.

MaTERIAL UXAMINED.—SOLOMON iSLANDS:

Kolombangara: Corner RSS 1188, 5500 ft., st., 2 Sep.

1965 (K); Guadalcanai. Corner RSS 89,

Popomanasiu, 7000 ft., fl., 23 Oct. 1965 (K, L); Hill

RSS 9011, Mt. Popomanasiu, 7000 ft., st., 5 Nov.

1965 (K, L).

LOCAL NAME.—Huta-ana-kwai ( Corner RSS

89).

The description is based on Corner RSS 89, the

only fertile collection, because stérile materiai of

Weinmannia is notoriously variable and often

lacks the distinguishing characters seen in adult

plants.

This species belongs to the same group as

Weinmannia croftii from the Bismarck

Archipelago and Karkar Island, W. denhamii

front Vanuatu, and W. vitiensis from Fiji, since

ail four taxa hâve a winged petiole and rachis,

and an inflorescence consisting of triads or pen¬

tads, as trequenfly see in sect. Leiospermum. It

may in fact turn out to be conspecific with W.

ctoftii, but more fertile collections are needed to

déterminé whether the range in leaflet size and

shape for this population is unique.

6. Weinmannia sp. (Solomons B)

Parker 6, from Mt. Bolbi, Bougainville Island,

Papua New Guinea, in low forest 10-30 fi. high,

8000 ft., Oct. 1963 (CANB) is a specinten in bud

only of an undescribed species belonging to sect.

Leiospermum.

ADANSONIA, sér, 3 • 1998 • 20(1)

Weinmannia in New Guinea and the western Pacific

Young stems hirsute, branching sometimes

dichotomous. Stipules not seen. Leaves impari-

pinnate, with 3-5 pairs ofleaflets, up to ca. 5 cm

long: pétiole and rachis segments 0.4-0.7 cm,

hirsute especially on upper surface, winged, the

wings extending up to 0.7 mm on either side of

the midline; latéral leaflets ± elliptical, 0.7-1.3 X

0.4-0.5 cm, base acute, apex broadly acute; ter¬

minal leaflets elliptical, 1.5-1.9 X 0.5-0.6 cm,

base attenuate, apex acute; blades coriaceous,

glabrous. Floral buds inserted singly on inflor¬

escence axis.—Fig. 5.

III. WEINMANNIA IN VANUATU (with

R.D. Hoogland )

Two species occur in this région, both of them

endemic: Weinmannia denhamii is widespread

and more common than W. JndcgilUprayi. The

first belongs to the grôup of species in sect.

l.emspemium that hâve a narrowly winged leaf

rachis and are often found on young volcanic

soils, where as U7 macgillivntyi is closely allied to

W. firaxinea in sect. Fasciculata. Both species in

Vanuatu hâve predominantly unisexual flowers

and almost glabrous fruits.

Key to the species of Vanuatu

1. Leaf rachis and petiole terert, densely haïr)' on the upper surface, not winged; latéral leaflets often with

small “cars” at point of attachmcnt to leaf rachis; inflorescence of one or more pairs of latéral dyads, the

apical bud of main stem persisting and dormant during flciwering .1. W. tnacgillivruyi

1’. Leaf rachis and petiole semiterete, glabrous, winged; latéral leaflets without “ears" at point of attachaient

to leaf rachis; inflorescence a central triad or pentad sometimes also with latéral triads or addttional

racemes .2. W. denhamii

1. Weinmannia maegillivrayi Seem.

Fl. Vit.: 109 (1866); Engl-, Nat. Pflanzenfam., ed.

2, 18a: 256 (1930); Bernardi, Bot. Jahrb. Syst. 83:

173, t. 22 (1964).—Type: Milne 277 (Vanuatu),

Aneiteum (= Anatotn), H.M.S. Herald 1854 (lecto-,

here designated, K!).

Small shrub to largish tree, 15-20 m tall.

Branching not usually dichotomous. Nodes

somewhat laterally flattened, the stipulât scar

arched on both sides, especially at distal nodes.

Stems glabrescent, rarely not, with numerous

lenticels and fine longitudinal grooves. Stipules

usually caducous, ± orbicular or shortly sparhul-

ate, roundcd at apex rounded and narrowing

towards base, up to 1 X 1.1 cm, sometimes revol-

ute, shortly strigose on abaxial surface, especially

towards the base and glabrous on adaxial surface

except at base. Leaves imparipinnate with 4-10

(-12) pairs ofleaflets, up to 14 cm long inclu-

ding petiole of 10-35 mm; rachis segments 5-

9 mm long; petiole and rachis terete or

channelled, densely hatry on adaxial surface and

in groove, the hairs up to 0.5 mm long, glabrous

or with few scattered hairs below (rarely densely

hairy ail round, Veillon 3992); latéral leaflets ses-

sile, narrowly ovate to lanceolate, the lower 1-3

pairs disrinçtly shorter than the upper pairs but

of similar width, up to 2.2-4.6 x 0.6-1.3 cm,

apex narrowly acuminate, base unequal, obtuse

to roundcd, not iying fiat on pressing, often 1 or

2 small frec auricles ca. 0,6 x 0,4 mm on rachis

between bases of opposite leaflets; apical leaflets

2.7-5 5 x 0,9-1.4 cm, narrowly elliptical, the

base attenuate, the apex narrowly acuminate to

caudate; blades glabrous on both surfaces except

for weak hairs ca. 1 mm long on rnidrib on

abaxial surface, not puncrate below, subcoriac-

cous; margin sometimes somewhat thickcned or

minutely revolute, crcnate to scrracc especially

towards the apex with up to 10 tecth on each

side of the latgest leaflets; rnidrib indented above

and prominent below, secondary veins ± fiat on

both surfaces.

Inflorescence of 1 or 2 pairs of dyads, some-

rimes a few successive nodes producing partial

inflorescences simultaneously; peduncles 0.2-

0.6 cm long, shortly and sparsely pubescent; api¬

cal bud of main stem between central peduncles

usually présent, sericeous; buds at apex of

ADANSONIA, sèr. 3 • 1998 ■ 20(1)

Hopkins H.C.F.

peduncles in angle bctween racemes serîceous,

minute; racemes ca. 7-14 cm long, rachis sparse-

ly puberulent. Floral buds inserred in fascicles;

floral bracts ligulate or somewhat boat-shaped,

0.4-0.9 mm long, eaducous with short, sciff

hairs. Flowers unisexual or rarely bisexual; pedi-

cel 1.6-2.8 mm long glabrous or minutcly

pubescenc at base; calyx lobes triangular, 0.6-0.7

x 0.4 mm long, glabrous; pétais elliptical-ovâte

or obovate, 1-1.6 X 0.7-0.8 mm; dise lobes 0.2-

0.4 mm long; in male flowers: filaments 2.8-

4.2 mm long, ovary smâ|l 0.3-0.9 mm,

pubescent, the styles 0.4-1.2 mm long and not

exserted beyond the antbers, in female flowers'

stamens short and not exserted beyond the styles,

the filaments up lo ca. l.I mm long, ovary ca.

1 mm long, pubescent, styles ca. 1.3 mm long,

ovules 8 per locule.

Capsules 2-3.5 X 2-2.2 mm at dehiscence, the

styles 1.7-2 mm long, the exocarp srnooth,

almost glabrous with a few weak hairs; calyx

lobes persistent, central column weakly develop-

ed or absent. Seeds ça. 0.8 mm long, persistent

in capsule, usually comose at both ends, the hairs

coarse and straw-like, ca. 2 mni long, twisted

around seed in capsule.—Fig. 7A-E, 8.

Breeding SYSTEM.—Polygamodioecious. Most

specimens bave only either male or female flow¬

ers but Kajewski 735 has male and a few bisexual

ones (though the ovary may not be functional as

the ovules are minute). In Schmid 3557 the flo¬

wers appear bisexual with a few males ones

mixed in.

FlELD CHARACTER.S.—Flowers white or créant,

fruits reddish.

Distribution and ecolocy. —Several collect¬

ions from Anatom (also known as Aneityum),

the Southern most of the main islands in the

chain, and one from Sanco, the largest island,

towards the north. Recorded from 30-300 m

from scrub, tain forest. open secondary forest

and forest on ridge. Described as comrnon

{Kajewski 735).

Material examined.— Vanuatu: Anatom

(Aneityum)-. Bernardi 12974 , nr. Anawounamalo, R.

Inwa Lelgey (sic) (= Imvan Lclcghci), 10-180 m, st., 5

May 1968 (K, P); Bourdy 322 , autour d’Anelgahoat

(sic), 30 m, fr., 1 Feb. 1986 (K, P); Cabalion 1944 , W

Inwoouné R., 300 ni. fl., 18 Mar. 1983 (NOU, P);

Cabalion 1944b, (Anelgaoh.it, Inwoourié R., Res.

Service Forestier, 80 m, lr., 18 Mar. 1983 (NOIJ, P);

Kajewski 735 , Anelgauhat Bay (sic), 100 fr., buds, 11

Feb. 1929 (A, RISH. K. P); R a ata 14. fr., 1976

(NOU); Schmid 3557. fl., 1 Dec. 1970 (P); Schmid

5083. secteur de Anelghanal, fl., fr., Nov. 1974

(NOU); S.coll.. s.n.. s.loc. (BM, type folder, no label).

Espiritu Santo: Veillon 3992, direction Voutmele,

buds, Aug. 1979 (P).

LOCAL N AM LS. —Nomropon ( Bourdy 322) but

see under Weinmannia denhamii in WHEATLEY

(1992).

TYPIFICATION.—The material of both this spe-

cies and of Weinmannia denhamii described by

SEEMANN was collected during the voyage of the

H.M.S. HltRALD, under the command of Câptain

H.M. DenhAM. MACGu.1.1 VRAY wàs the ships

naturalist and Mil.NE the assistant n.uuralist befo-

re they were dismissed and discharged rcspectively

during the voyage, for dcreliction of duty (David

1995). According to DAVID, the Herald visited

Aneiteum, where the collections of Weinmannia

were made, between 7 November 1853 and

1 January 1854, and plant specimens were sent to

W. HOOKER at Kew However, Mac.Gillivray

may hâve passed sonie of Mir.NF.'s collections off

as his own, and 11OQ KL R criricised the pour Iabel-

ling of the material he received (David 1995). In

the description of W. macgjUivrayi, Sf.EMANN cited

material collected by both MacGili ivjlay and

MlLNL, which constitute syntypes; Milne 277 (K)

is designated here as the lecrotype. A specimen in

a type folder at BM has no label and may or may

noc belong ro the same collection.

ReLA'I'IONSHIPS.—T his spccics is a satellite of

the widespread Malesian W. jraxinea, distingui-

shed from it by more numérotas and smaller leaf-

lets and by the cars that axe frequent on the leaf

rachis at the point of insertion of the latéral leaf-

lets. In both species, the upper surface of the leaf

rachis is often densely hairy.

2. Weinmannia denhamii Seem.

Fl. Vit.: 109 (1866), as “denhami”; Engler, Nat.

Pflanzenfam., ed. 2, 18a: 256 (1930); Bernardi, Bot.

ADANSONIA, sér, 3 • 1998 • 20(1)

Weinmannia in New Guinea and the western Pacific

Fig. 7.— A-E. Weinmannia mscglllivrayi: A. part of a îlowertng shoot with impanpinnate leaves (only one shownl at the most distal

node subtendina an Inflorescence ot 4 dyads; more proximal node has a latéral shoot with an inllorescence of two dyads: note minu¬

te buds in angle between racemes of each dyad; B, flower, probably biseyuaf: C. lemale flower, one pelai removed; D, dohisced

capsule with persistent calyx lobes E t seed. {A, Ka/ewsM 735: B, Schmid 3557 ; C, Cabalion 1944, D, E, Roata 14), —F-N,

Weinmannia denhamii F part o» a tlowenng shoot with central and latéral triaOs (only one latéral triad shown), note reduced leaves

at some nodes within inflorescence. G, single leal with smaller, more numerous, leaflets (type of W tannaensis), H, mate flower; J,

detail of H with perianth removed to show gynoecium with very reduced styles, dise lobes and a single stamen; K. female flower; L,

detail of K wilh perianth removed to show gynoecium with long styles, dise lobes and a single stamen, M, capsule at dehiscence,

note central column and enlarged réceptacle; N, seed. (F, MacGillivray 45: G, Kajewski 151: H. J, M, N, Aubert De La Rue s.n.,

2ème Voyage, Jan. 1936; K, Schmid 5084). —Drawn by C. Monnié.

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

Jahrb. Sysi, 83: 190, t, 32 (1964); Wheatley, Guide

Commun Trees Vanuatu: 76, t. 77 (1992).—Type:

(. MacGillivray ) 45, (Vanuatu), Aneiteum (= Anatom),

in woods dette-, here designated. K!; isolecto-, BM!,

G, P! 3 sbects).

Weinnuinnia kajewikit Guillaumin. J. Arnold Arbor.

12: 250 (1931).—Type: Kajewski 317 (Vanuatu),

Eromatiga Island, Dillon Bay, 400 m, rain forest,

commoD, 28 May 1928 (holo-, P 1 ; iso-. A!, BTSH1).

Weinmannia wnnaetisis Guillaumin, ), Arnold Arbor.

12: 251 (1931); Bernardi. Bot. Jahrb. Syst. 83: 178,

t. 27 (1964).- Type: Kajewski 151 (Vanuatu),

Tanna Island, Mr. Tokosh Mtru, 1000 m, rain

forest, commun, 15 Mar. 1928 (liolo-, P!; iso-,

BISH!).

Weinmamùa kajewskii Guillaumin var. ambrymensh

Guillaumin, Bull. Soc. Bot. France 82: 349 (1935),

nom. illeg., no Latin description. Collection cited:

Aubert de la Rite s. n. (Vanuatu), Ile Ambryrn, sur le

grande plateau de scories à proximité du volcan

actif, cote 600-700, Août 1934 (A!, L!, P!).

Shrub or tree, 2-20 m high, also sometimes an

epiphyte or srrangler; hranching sometimes

dichotomous; young twigs shortly hairy, finelv

ridged with narrow longitudinal fissures and

round or elliptical lenticels; older rwigs with stip-

ular scars ± horizontal Stipules usually caduc-

ous on reproductive branches, sometimes

persistent on végétative ones, variable in shape,

usually ovare or ligulate, 6-10 x 2-4.5 mm,

sometimes orbicular (juvénile loliage?), 8-12 x

10-14 mm, rounded or obtuse at apex, glabrous

or minutely hairy on adaxial surface, with a few

scattered hairs to 0,3 mm long on abaxial surf¬

ace. Leaves imparipinnate with 1-12 pairs of Icaf-

lets, up to 14 cm long iricluding petiole of

7-25 mm; rachis segments 6-13 mm long; petio¬

le and rachis segments semiterete, flattened above

with a narrow central ridge and winged, the

wings extending 0.5-1.5 mm on cicher sidc of

mid-line, on eaeh segment the wings broader

distally towards point of attachaient of leaflets;

petiole and rachis glabrous to pulvérulent above,

glabrous or bearing a few hairs on underside

especially at point of attachaient of leaflets; latér¬

al leaflets lanceolate, elliptical or ovatc, the prox¬

imal ones often shorter rban the more distal

ones, l.l-3(-4.8) x 0.3-1.1(-1.5) cm, apex acute,

base asymmetrical, distal side acute to attenuate,

proximal side obtuse ro rounded; terminal leailet

narrowly elliptical to narrowly obovate 2.1-4.1

(-6.4) X 0.3-1 (-1.7) cm, apex acute, base attenua¬

te; in leaves with few pairs of pinnae, terminal

leailet often larger than latéral ones; in leaves

with numerous pairs of pinnae, terminal leaflet ±

equal length and uarrower than latéral ones;

blade glabrous cm both surfaces, usually puiictate

below, subcoriaceotis; margin somewhat thicken-

ed and minutely revolute, crenate especially

towards the apex with 6-17 crenations on each

side; midrib narrowly prominent above and

slightly prominent below, when sometimes

minutely hairy.

Inflorescence a central triad or usually pentad,

the lower racemes either in the axils of leaves or

not, rarely replaced by latéral triads; sometimes a

few successive nodes on a shoot bearing partial

inflorescences simultaneously; peduncles and

rachis segments 0.8-3 cm long; racemc-s 5-

11.5 cm long; inflorescences axes puberulent.

Floral buds inserted singly; floral braers lanceo-

lare or somewbat carinare, up to 1.1 mm long,

bearing a few hairs towards base on abaxial surfa¬

ce, caducotts. Flowers unisexuai; pedicel ( 1 -) 1.4-

3 mm long, slender, puberulent: calyx lobes

triangular, 0.9 x 0.6-0,7 mm, ± glabrous; petals

ovate-ellipdcal, 1.3-1.6 X 0.7-0.9 mm; dise lobes

0.5-0.7 mm long; in male flowers: stamens long

and anthers far exserted beyond stigmas, fila¬

ments ( 1,6-)2-3,5 mm long, ovary 0.9-1 1 mm

long, styles minute, < 0.1 mm long and curved

inwards; in femalc flowers; stamens short and

Stigmas exserted beyond anthers, filaments 1-

1.7 mm long, ovary 1.3-1.5 mm long, almost

glabrous, sryies ca. 1 mm long.

Capsules distinctly supporred by réceptacle,

valves 3-4 X 1.7-2 mm at dehiscence, the styles

0.8-1.3 mm long; exocarp with minute longitud¬

inal ridges, almost glabrous with a few minute

strigose hairs; calyx lobes not persistent; central

column often persistent on réceptacle after valves

hâve fallen. Seeds ca. 0.6 mm long, 16 per capsu¬

le, tending to persist in capsule after dehiscence,

cornose at both ends, the hairs 0.8-1 mm

long.—Fig. 7F-N, 8.

BREEDING SYSTEM.—Dioecious or polygamo-

dioecious? Most specimens bave unisexuai flow¬

ers bur two collections at P {Aubert de la Rite s.n.,

19 Feb. 1936 and Bernardi 12986) hâve both

ADANSONIA. sér. 3 • 1998 ■ 20(1)

Hopkins H.C.F.

male flowers and fruits on rhe saine twig.

GROW'IH FORM AND FIF.tD CHARACTERS. —As

in other species whose distribution covers a wide

range of élévation, size and grovvth habit are

influenced by alrirude and exposure. It is record-

ed as a tree 20 m high at 1300 m (on Ambae)

and as a shrub 2 m high at 1800 m (on Santo),

When a large tree, rhe crown rnay be open and

spreading, flar-topped with che branches vertical-

ly ascending to form an in vert ed cône, or donte-

shaped vvith divaricate branches, the fotiage

feathery (Wbeatley 207), in distinct clumps on

otherwise bare branches ( Wbeatley 55). The bole

is sometimes twisred and leaning. Also described

by Wbeatley 207 (and 1992) as a strangler that

kills its host and then forms a bole by the fusion

of the roots, with a diamerer of nearly 70 cm.

Bark light grey or brown-grey, smooth with fine

vertical fissures, blackish or reddish on old

trunks Inner bark red or orange, fibrous. Sap

wood wbite, cream to yellowish. Heartwood

pink-cream, sweet-smelling; Young leaves flush

reddish or ycilowish-red, mature leaves dark

green and sJightly shiny above. silver-green below

{Wbeatley 207). Flowers usually described as

white, rarely yeilowish-green or pinlt, and much

visited by insects ( Bernürdi 12986). Fruit pink,

red then brown.

Distribution and ecology. —Known from

most of the larger islands of Vanuatu. Lowest

recorded altitude varies by island but the overall

élévation range is 120-1800 m.

According to WHEATLEY (1992), W. denhamii

is a canopy species of upper middte and high

élévation lorest where it is co-dominant with

Metrosideros over large areas. Other habitats

include riverine forest on basait, open woodland

with tree ferns, bracken-covered ridge grazed by

bullocks, open Acacia spirorbts scrub, open

secondaty lorest, and deforested areas with

Pteridiurn and Gleichenia. At higher élévations it

occurs in montane, dwarf foxest on exposed

crests and ridgcs.

Soil types recorded trom speeimens include vol-

canic cinders and lava, and young brown ando-

sol. Aithough W. denhamii belongs to the group

of species (including W. crofiii , W. vitiensis and

W. sp. Solomons A) that are often found on rela-

tively young volcanic soils, WHEATLEY says it is

less frequent than Metrosideros on the juvénile

ash soils of Tanna and theAmbrym caldera.

Material examined. — Vanuatu: Vanna Lava.

Wbeatley375, s.loc., 550 m, fl. (K). (Espiritu) Santo:

Cabalion 462, Col sur le Custom rd., entre R. Pialapa

et village Wunabaï, 850 m, fl., 21 Mar. 1978 (NOU);

Cheiv Wee-Lek RSNH 204, Mt. Tabwetnasana,

1520 tn. st., 3-8 Sep. 1971 (K); Cribb de Wbeatley 82,

Saktouy to Tabwemasana, 1500-1550 ni, fr., 24 Oct.

1988 (K); Curry I640B, Fortsenale, nr, village,

700 m, fr., 10 Dec. 1994 (K); Gillistm dr fteveridge

RSNH 3531, Nukowoula village area, 3500 ft., fr,, 28

Aug. 1971 (P): Raynal RSNH 16345, Mt.

Tabwemasana. 1800 m, ft., 2 Sep. 1971 (K, L);

Veîllon 4026, direction du Vommelc, 1000 m, fr.,

Aug. 1979 (P). Ambae (= Aobd): Quart tin in Sebmid

1041, sommet du volcan, 1000 m, st., 18 Nov. 1966

(P); Wbeatley 55, s.loc., 1300 m, st., 23 Nov. 1988

(K). Penteeort: Aubert de le Rite s,n, Kumrv, 600 m,

st., 19 Dec. 1935 (P); Wbeatley 207, Tambok, 460 m,

buds, 9 Feb. 1989 (K). Ambrym: Aubert de le Riie s.n.,

Mt Folio, 1000 m, st., 5 Jan. 1936 (P); Aubert de le

Rite en., 650 800 m, fl., fr., 8-11 Jan. 1936 (F, P);

Aubert de le Rite tn.. Grand plateau autour du Mt.

Marutn. 700 m, fl., fr., 10 fan 1936 (P); Bourdy 118,

au-dessus de Lalinda, 600 m, fr.,. 23 Sep. 1985 (K, P).

Epi. Curry 1026, Mt. Tainaruru, close to summit, S

sidc above Filakara, 500 m, fr., 20 july 1993 (K).

Efate: Sebmid 248, Vate, 450 m, fr., 26 Feb. 1965

(P); Kajemki 236, Undine Bay, 500 m. fr., 25 Apr.

1928 (K, P). Erromango: Aubert de le Rite s.n., W du

massif occupant le N de F fie, 400 m, fl., h.. 19 Feb.

1936 (P); Bernardi 13258, W. Di lion Ilay,

Oponhkor. Mt. Mcnel, R. Nevel, 250-350 m. fr., 3

lune 1968 (K, I„ P); Cabalion 1388, S, l edmoghum,

300 m, fr., 21 Feb. 1982 ( N O F) ; Chtesnutn 34, s.loc.,

1600 ft, h,, 17 July 1930 (K); Ciory 350, Fogging

trail from DiUons Bay to Port Marvin, 120 m, ir., 5

Aug, 1992 (K): Johnson 12, s.loc., 300 m, fl., s.dat.

(K); Sebmid f740. Centre, 300 m, buds, fr., 25 June

1973 (NOU); Sebmid 308a. N de Happy I.and,

250 m, fl, 21 Nov. 1974 (NOU). Tanna: Aubert de

le Riie s.n., s.loc., st., Mar. 1934 (P); Berndrdi 12849,

nr. Fooropounga, 250-300 m, buds, 29-30 Apr, 1968

(K, F, P); Bernardt 12009, Lamwinaoura, fl„ b. 1-2

May 1968 (K, F. P); Bmtardi 13097, Paei Yanehoop,

300 m, fl., fr., 20 May 1968 (K, F, P); Chrw Wee-Lek

RSNH 109, Yeruu, SW font of Mt. Toukousmerou,

100-200 m, fl., fr„ 27 July 1971 (K, P); Morar 5897,

Flan E du Toukosmereu. 500 m, fl.. 7 Feb. 1978

(NOU, P); Moral 6043. Mt. Toukosmereu, 500-

600 m, fr., June 1978 (NOU. P); Sebmid 3176, vers

Côte Ust. 500 ni. ft.. 7 May 1970 (K, NOU, P).

Anatom (Aneityum): Bernardt 12927, nr.

Anawounamalo to R. Inwa Felgey, 10-180 m, fl., yfr.,

5 May 1968 (K, F, P); Bernardt 12986, nr.

Anowounamalo towards Anitchavo (= Port Patrick),

Ougaphaérek, lntéro, Ounétchnytap, to R. Inwa

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in New Guinea and the western Pacific

Anetcho, 430 m?, fl., £r., 7-8 May 1968 (K, L, P);

Bernardi 13018, ibid., 700 m, buis, 7-8 May 1968

(K, L, P); Mormon s.n., s.loc,, fr., 26 June 1896 (K);

Schmid3558, Sud, buds, Dec. 1970 (NOU).

LOCAI NAMES. —According to WHEATLËY

(1992) names refer ro die genus in general, e.g.

Nomropom (Aneiteum), Nariop (Erromango),

Aisensu (Paania), Kanumtep (Pentecost).

Additional names froni spécimen labels include:

Narurururu (Efate), Owara (Fortsenale language,

Santo), Nivlewi (Sie language, Erromango,

meaning: drinking medicîne [Curry 550\),

Sasatré (Santo).

Locai USES. — flic bark gives a red dye used on

clubs ( Aubert de la Rik s.n., 19 Dec. 1935). The

wood is hard and durable and used for “hoc” ftre-

wood and bouse construction (e.g. bouse posts

and rafters) (WHEATLEY 1992; Wheutley 207). It

is also used as medicine for asthma or “short

wind” (Curry 550)-. the bark is srripped off and

ground ro a powder which is then put into a

curled Heliconia leaf and cold water added; the

leafis squeezed and the extract drunk.

Typification.—Seemann gave rhe type as

“-Aneiteum, in woods (M'Gillimiy!)' . Sheets at

K and BM lack a collectons name but beat the

number 45. According to STAFLEU & COWAN

(1985), Sf.F.MANNs study set for the botany ot

the H.M.S. HERALD is at BM, but he publfehed

the names Weinmannia denhamii and W. mac-

gillivrayi in his “Flora Vitiensis”, for which K,

and not BM, is considered to hâve the first set.

Since the situation is somewhat confused, a

lectotype is designated at K. The isolectotypes at

P give both MacGillivRAYs name and the

number 45. The date “Feb. 1859” is after

MaCGili IVRAY had left the H.M.S. Herald

(David 1995), and may refer either to when the

material was received in London or when it was

collected on a later expédition (sec DAVID 1995:

218).

TaXONOMY AND VARIA I ION. —The leaves of W.

denhamii are variable in both the number of

pairs of leaflets and their shape. When there are

few leaflets, they are usually lanceolate and when

they are numerous, they arc shorter and relatively

broader. Flowever, there is no discontinuity in

the number of pairs of leaflets per leaf (R.D.

HOOCJLAND, unpublished, based on analysis of

38 sheets). Specimens with fewest leaflets are

from Efàtê, Santo and Anatom, and those with

the rnosrare ffom Tanna and Arnbae.

BERNARDI (1964) placed Weinmannia tannaen-

sis in sect. Fasiculata but the flowers are inserted

singly and the inflorescence architecture is thut of

sect. Leiospermum.

IV. WEINMANNIA IN FIJI

(with J.C, Bradford)

We recognize four species occurring in Fiji. In

sect. Fasciculata , Weinmannia richii is endemic

and relatively abundant on the rwo largest islands

while W. exigua. is known from Vanua Levu and

also occurs in the Solomon Islands. In sect.

Leiospermum , W. vitiensis is endemic and known

only from rhree of the smaller islands and W.

affinis is relatively abundant on Viii Levu,

Taveuni and Ovalau, and has also been reported

from Samoa although its status there is uncertain.

Key to the species in Fiji

1. Inflorescence a sériés of dyads or tetrads, the apical bud of the main stem présent between the peduncles

of the inner most partial inflorescences; latéral, auxiliary buds absent.2

1’. Inflorescence of triitds or pentads, clic apical bud of the main stem either dcveloped into a partial inflores¬

cence or aborted, latéral auxiliary buds présent .....3

2. Leaves on reproductive shoots usually triloliolate. the terminal leaflet often mucli larger rhan the laterals ..

.... 1. W. richii

2’. Leaves on reproductive shoots usually imparipinnate with (l-)3-7 pairs ofleaflets, the terminal leaflet not

or scarcely larger than the largest laterals .. 2. W. exigua

3. Leaves on reproductive shoots unifoliolate.3. W. arnnis

3’. Leaves on reproductive shoots usually trifoliolate, the rachis narrowly winged .4. W. vitiensis

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

1. Weinmannia richii A. Gray

U.S. Expi. Exped., l’han. 1: 675, Atlas t. 85 (1854);

Seem., Fl. Vit.: 110 (1865); Engl., Linnaea 36: 643

(1870), Nat. Pflanzenfam., ed. 2. 18a: 255 (1930);

A.C. Stn., J, Arnold Arbor. 33: 133 (1952); Bernardi,

Bot. Jahrb. Syst. 83. 177, t. 26 (1964); J. Parliam, PI.

Fiji Islands: 124 (1972, rev. etL); A.C. Sm„ Fl. Vit.

Nova 3: 20, fig. 5D (1985).—Type: US 48071, Fiji

(Feejee) Islands, on mountains ar SandaKvood Bay

[= Mbua Ba y ficie Smith), Vanua Lcvu, 2000 (t., fr.,

1840, US F.xpl. Exped. undet the command of’Gapt.

Wilkes (holo-. US 48071!; iso-, BM!, GH, K!, NY).

Weinmannia rhodagyne Gibbs, J. Linn. Soc. Bot. 39:

145 (1909); Engl., Nat, Pflanzenfam., ed. 2, 18a:

255 (1930).—Type: Gibbs 504, Fiji, Viti Lcvu,

Nandàrivatu, 2700 ft., fl., Aug 1907 (holo-, BM!;

iso-, K!).

Shrub or small tree 2-10(-20) in high.

Branching not usually dichotomous. Nodes

somewhat laterally flattened, che stipular scar

arched on both sides, especially at distal nodes;

internodes ca. 0.8-4.5(-8.5) cm. Young stems

puberulent, older ones glabrous wirh numerous

lenticels. Stipules caducous or not, suborbicular,

up to 1 1 x 1.5 cm, rounded at apex, strigose ro

glabrous on abaxial surface. I caves usually com-

pound (1-)3(-7)-foliolate; total length up to

11 cm (for a trifoliolate leaf); petiole 0,9-1.5 cm

long in trifoliolate leaves, to ca. 3 cm in impari-

pinnate ones; petiole and rachis segments where

présent subterete, slightly flattened or channelled

on adaxial side, especially distally towards point

of insertion of leaflets, scaly or puberulent;:

leaflets elliptical, rarely narrowly elliptical, lan-

ceolate or ovate, often conduplicate when dry; in

a trifoliolate leaf, latéral leaflets 2.1-5 X 0.7-

3.9 cm, sessile, base t equal to unequal, apex

acute; apical leaflet often much larger than the

laterals of the same leaf, 3.3-8.2(-9.5) X 1.1-3.2

(-4.4) cm, base attenuate, apex acute; in impari-

pinnare leaves, largest laterals about equal to

terminal leaflet but not attenuate at base; blades

subcoriaceous, glabrous, nor punctate below;

margin sometimes thickened, somewhat sinuate-

crenate and unduiate, ca. 8-12 crenations on

each side of a leaflet; midrib slightly depressed

above, prominent below and sparsely hairy,

secondary and tertiary venation slightly raised on

both surfaces.

Inflorescence usually (2-)4(-6) dyads or terrads

in a sériés at a node, sometimes a few successive

nodes producing dyads or tetrads; peduncles 0.5-

12 mm long, shordy strigose or with short erect

hairs or almost glabrous; buds at most distal

node of main stem and between che central pair

of peduncles 1 or 3, sericeous; buds at apex of

peduncle in angle between central pair of

racemes sericeous, minute; rachises shortly

pubescent, up to 10 cm long. Floral buds insert-

ed in fascicles, tending to dissociate from the

bract, the fascicles somewhat verricillate and the

vertteils somewhat separared so that sections of

bare rachis visible between chem; floral bracts

about equal in length to buds, L5(-2) mm long,

cymbifoim, caducous, with shorr stiff hairs.

Flowers unisexual or bisexual; pcdicel 1-1.5 mm

long, almost glabrous: calyx lobes 0.5-0.6 mm

long, glabrous; petals obovatc (not oblong) and

often retuse ai apex, 1-1.3 X 0.7-0.9 mm; dise

lobes ca. 0.3 mm long; in female flowers: fila¬

ments 1-1.2 mm long, ovary pilose to densely

pubescent, ca. 0.9 mm long, the styles ca.

0.9 mm long, ± straight or curved outwards; in

male and bisexual flowers: filaments up to 2.2-

3.8 mm, anthers ca. 0.4 mm long; in male

flowers: ovary 0.4-0.5 mm long, the styles 0.4-

0 5 mm long, curled inwards: in bisexual flowers

(post anthesis): ovary ca. 1.1 mm long, the styles

1 1 mm long, straight or curved outwards.

Capsule 2-3 5 x 1.2-2 mm at dehiscence, the

styles 0.5-1.5 mm, the exocarp almost smooth,

softly pubescent; calyx lobes usually persistent;

central column weakly developed, usually not

persistent after valves hâve fallen. Seeds 0.8-

1 mm long, 8 per capsule, comose at each end,

the hairs to 1.5-2 mm long, much curled in cap¬

sule, later straight, sometimes with shorter hairs

over the rest of the testa.—Fig. 9D-J, 10.

JUVENILE FOLIACE. —Hopkins dr Bradford 5024,

sucker shoot in dense shade: leaves with up to 6

pairs of pinnae; leaflets chartaceous with serrate

margins; stipules persistent, rounded and salver-

form, forming a horizontally flattened ring

round the stem at nodes.

BkFFDINC SYSTEM. —Polygamodioecious.

FlELD CHÀRACTERS. —Shrub or small, compact

tree with a dense, rounded or irregular crown.

ADANSONIA, sér. 3 • 1998 ■ 20(1)

Hopkins H.C.F.

Flowers whitc, grccnish white, creamy or some-

what pink and the pedicel pinkish; also recorded

as yellow orange (TURRILL 1915).

Distribution and ecology. —Known from

Viti Lcvu and Vanua Levu. Reported by

BERNARD) from Samoa on the basis of stérile

collections vvhich are unlikely to be this species.

Occurring Irom 100-200 up to 1400 m. with

the majoriiy ol records from 800-1000 m.

Habitats include varions types of forest (montane

forest, ridge forest, open and dense forest [with

tall gingers and tree ferns]) on ridges, slopes and

in rolling country; also scrub and thickets in

grassland and cm dry slopes. Sometimes locally

comnton. Field observations in central Viti Levu

in March 1996 found that adult trees were often

clumped and ntost were in fruit. There was a

high degree of phenological symehrony both

within individual trees and within populations

(pers. obs.).

Selected collections (from a total of 32 stud-

ied).—Fiji: Viti Levu: Damante FD 991 D.177,

Serua, Nabuutinî, 1000 lu, buds, 21 |att. 1964

(SUVA); Degettn 14379, Thohi North, nr.

Nandarivatu, Nandala fish hatchery. 750-900 ni, ft..

4 l ; eb.-26 Mar. 1941 (A, BISH, K, L, P); Gillespie

4233, Tholo North prov.. Nardarh'atu, 2 miles along

Nandran trail, 850 m. fl., 8 Dec. 1927 (BISH, GH);

Gillespie 4333 , Tholo North prov,, slopes of Loma

Langa Ml-, 1 100 m, fl.. fr„ 19 Dec. 1927 (BISH,

GH); Greenwood 384, Luutoka, 600 ft., lr.. 23 Oct,

1931 (A, K); nr. MonasavU Dam, I km N oIFLA sta¬

tion, 840 m, fr., 28 Mar, 1996, Hopkins & Bradford

5023 (MO, P. SUVA); im Thurn 73, Nandarivatu, (1.,

1911 (BM, K, P); Koroiveihau DA II 129, Nailaga,

Ba, buds, 19 Dec. 195” (K, SUVA); Kuruooli DA

16061 , Macuata, Mt. Delainacau, fl.. 9 Dec. 1968 (A,

BISH, P. SUVA); Parbam DA 7121, berween Navai

and Nad.trivatu, Mba. Prov. Ba, 4340 ft., buds, fr., 15

Feb. 1951 (BISH, SUVA); Qtiro DA 12947, Macuata,

Tadradave, buds. 5 Nov. 1968 (SUVA); Ranamu

(DA) FD 1174 , Nadrao, Navosa, Nabosewale,

Nadrau, 2700 ft., H., 19 May 1967 (BISH. SUVA);

Smith 4710. Nandronga, Navosa (fcuinerly Tholo

West), S slopes Nausori Flighlands, Namosi Creek

above J umbenasolo, 300-450 m, fl., 29 May-5 J une

1947 (A, BISH, !.. P); Smith 5739, Mba, fdrmerly

Tholo North, W slopes of Mt. Nanggaranambuluta

(Lomalangi), £ of Nandarivatu, 850-1000 m. fr., 19

June-2 Oct. 1947 (A, BISH, K. L. P). Vanua Levu:

Parham DA 1122, Bua or Mbua, Wairiki, 11., I May

1938 (A, SLIVA); Parham & Seitlo/o DA 2277, Bua or

Mbua, Ndama, 90 m, fr., 7 Dec. 1939 (A, SUVA);

Parham s.n., Bua Prov., fl., Jan. 1937 (A, BM); Smith

6813, Mathuata, Seanggangga Plateau, Korovuli R., nr.

Nanta, 100-200 m, fr., 25 Nov.-8 Dec. 1947 (K, L, P).

Local namêS.—Q alo (Parham 1941, sub W.

rhodogyne)\ Manaui (Parham DA 1122 ); Vota

(Smith 6813).

2. Weinmannia exigua A.C. Sm.

J. Arnold Arbor. 33: 137 (1952); Bernardi, Bot.

Jahrb, Syst. 83: 166 (1964); J. Parham, PI. Fiji

Islands; 122 (1972, rev. ed.); A.C. Sm., Fl. Vit. Nova

3: 25, Fig. 5C (1985). —Type: Honte 632, Fiji, Vanua

Levu, berween Waiwai and Lomaloma, May 1878, fl.,

yfr. (holo-, K!).

Shrub or small tree, ca. 3-13 m high. Branching

not usually dichotomous. Stems woody, not

thickened at nodes and annular scar faim, inter-

nodes 0.5-2.3 cm long. Young stems minutely

hatry, axillary buds dcnsc-ly hirsute, older stems

glabrescent, bearing numeroUs pale lenticels.

Stipules mostly caducous even in végétative

ntaterial, rarely seen, t orbicular in outline with

a narrowed base, ca. 0.5 X 0.5 cm, strigose on

abaxial surface especially towards the base and

glabrous on ad axial surface, margin somewhat

recurved. Leaves usually imparipinnate, (l-)2-9

pairs ol latéral leaflets, rarely unifoliolate, total

length up to 5 cm long; petioie and rachis seg¬

ments semiterete and very narrowly winged, the

wings sometimes curved upwards to give a later-

ally compressed U-shaped cross section, often

densely pubescent on adaxial surface along

midrib, more sparsely hairy elsewhere, petioie up

to 1 cm long (0,3 cm in unifoliolate leaves),

rachis segments 0.3-0.5 cm long; latéral leaflets

elliptical or sometimes obovate, the distal ones

larger than the proximal ones (0.3-) 1-1.7 x (0.3-)

0.4-0.6 cm, hase acutc and ahnost equal, apex

acute; terminal leaflet narrowly elliptical, narrow¬

ly obovate to elliptical. 1.6-1 X Q.4-0.7 cm long,

base attenuate, apex acute; blades glabrous except

for midrib vvhich is sparsely srrigose below, coria-

ceous, not punctare; margin sometimes thicken¬

ed, crenulate with 4-6 notches on each side in

the latéral leaflets; midrib slightly depressed

above and prominent below, secondary veins fiat

or very slightly' raised above and below.

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in New Guinea and the western Pacific

Inflorescence a pair of dyads or individual

racemes, che apical bud of main stem densely

minute, velutinous; peduncles 0.8-1 cm long,

racemes up to 5 cm long; inflorescence axes spar-

velutinous, axillary buds at most distal leaf- sely hairy. Flowers inserted in fascicles; floral

bearing node olten présent and densely velutin- bracts caducous; female and bisexual flowers

ous; bud at apex of peduncles between racemes seen; pedicel ca. 1.5 mm long, minutely sparsely

Fig. 10.—Distribution of Weinmannia in Fiji

ADANSONIA, sér. 3 • 1998 ■ 20(1)

Hopkins H.C.F.

hairy; calyx lobes 0.6-0.7 mm long, glabrous;

petals 1.1-1.4 x 0.6-0.7 mm, rounded or nor-

ched at apex; dise lobes 0.4-0.5 mm long; in

female flosvers: lilamenrs ca. 1.3 mm and ovary

ca. 1.3 mm long (bue past anthesis), pubescent,

with styles 1 2 mm long; in bisexual flower.s; fila¬

ments ca. 4 mm long; ovary ca. 0 7 mm long at

anthesis. densely velutinous, die styles 1.7-2 mm

long. Immature fruit with ovary ca. 1.3 mm

long, densely golden velutinous and styles ca.

2 mm long, glabrous, the calyx lobes mostly fall-

en, dise lobes persistent. Mature capsules and

seeds not seen.—Fig. 5, 9A-C, 10.

Breeding SYSTEM —Polygamodioecious?

Home 632 has female flowers and Whitmore

BSIP 995 (from the Solomon Islands) has

bisexual ones.

FlELD CMARACTERS,—Habit appears to be

variable as the type is described as a large shrub

10 ft. high, while in the Solomons it is a large

tree.

Distribution and ecology. —Fiji (2 collec¬

tions seen) and the Solomon Islands (1 collection

seen). Habitats include high forest at 500-1000

ft. (Howard 89) and mountain top (Home 632).

MATF.RIA! F.XAM1NED.—Fiji: Vanua Leva:

Rangagone in Howard 89, Thakaundrove, nr. tributary

of Sovivi Crcek, S of Karoko, Tunuioa District,

Natew.i Peninsula, st., May 1968 (B1SH, K). Poorly

collectcd but apparently common in montane lorest.

Despite the wide disjunction, the leaves of the

collections from lhe Solomons and Fiji arc simil-

ar. Weinmannia urdanetensis also has numerous

small leaflets and is the species which most

resembles W. exigua but it dilTers in the details of

the shape, texture and venation of the leaflets,

and in the arrangement of the leaves. In the pro-

tologue, SMITH described the fruits and seeds but

there are none on the type at K.

3. Weinmannia affinis A. Gray

U.S. Expi. Exped., Phan. 1: 674 (1854); Seem., Fl.

Vit.: 110 (1865); Engl., Linnaea 36: 648 (1870), Nat.

Pflanzenfam., ed. 2, 18a: 256 (1930); Reinecke, Bot.

Jahrb. Syst. 25: 635 (1898); Gibbs, J. Linn. Soc. Bot.

39:145 (1909); Christoph., Bernice P. Bishop Mus.

Bull. 154: 9 (1938), p.p.; A.C. Sm., J. Arnold Arbor.

33: 130 (1952); Bernardi, Bot. Jahrb. Syst, 83; 187, t.

30 (1964); J. Parham, PI. Fiji Islands: 122 (1972, rev.

ed.); A.C. Sm., Fl. Vit. Nova 3: 19, fig. 5A,B,E

(1985).—Type: US 48070, Fiji (Fcejee) Islands, on

the mountains of Ovolau, 1200 ft., fr„ U.S. Expi.

Exped. under command of Capt. Wilkes (holo-, US

48070!; iso-, GH, K!, NY, P?).

Shrub or small compact tree 1-7 m high, rarely

larger (20 m) or an epiphyte. Branching sorne-

times dichotomous (or into 4), the apical bud

aborting, but not exclusively so; nodes somewhat

thickened; glabrous except for young stems,

inflorescence axes and pedicels shortly and spar-

sely strigose-pubescenr, axillary buds sericeous,

and ovaty sometimes with short, S par se indu-

mentum. Stipules variable in size, shape and per-

sistence: either rhombic, obcrullate, ovate or

broadly clliptical: either 0.5-1.1 X 0.35-0.6 cm

and usually caducous when leaves smali, or 1.1-

1.9 X 0.6-1.6 cm, coriaceous and persistent,

especially when leaves large; apex acute, almost

round to cuspidate; margin sometimes toothed.

Leaves simple, ohlong, elliptical or broadly ellip-

tical, the base decurrent into a short pétiole 0.4-

1.1 (-1.6) cm long, the bladc 1.8-9,8 x 1-4.8 cm,

the apex acute to obtuse; punctate below; margin

usually thickened, shallowly or markedly crenate

or rarely sinuate, 9-14 notches down each mar¬

gin of a leaf; midrib slightly raised above and

prominent below, secondary and tertiary vena¬

tion either Hat, raised or obscure above and

slightly raised below.

Inflorescence usually of 2 latéral triads or rarely

2 latéral pentads (central bud aborted), or some¬

times 3 triads (central bud developed); further

variants include four triads, developed from

basal, latéral auxiliary buds, the central bud abort¬

ed, a fcw successive nodes of the main stem pro-

ducing partial inflorescences simultaneously;

peduncles 0.7-2.7 cm long; racemes usually

short, 2.5-5 and not projecting beyond the folia-

ge, rarely to 9 cm long; stipules at nodes within

the inflorescence ligulate, sometimes persistent.

Floral buds inserted singly; floral bracts not far

excecding length of buds, ligulate to cvrnbiform,

ca. 0.7 cm long, not always lugaceous. Flowers

unisexual or bisexual; pedicel 0.9-1.8 mm long;

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in New Guinea and the western Pacific

calyx lobes 0.7-1.5 mm long; rarely pubescent;

pétais oblong or ovate, 1.2-1 .8 X 0.9 mm; dise

lobes 0.5-0.7 mm long; in male flowers: fila¬

ments up to 2.8-3 mm long, ovary ca. 1 mm

long plus styles of 0.2 mm; in female flowers:

filaments ca. 1 mm long; ovary (past anthesis) ca.

2 mm long plus styles 1.1 mm long.

Capsules 2.2-3.8 X 1.3-2.2 mm ai dehiscence,

the styles 1-1.3 mm long, the cxocarp somewhat

longitudinally ridged, glabrous orsparsely strigo-

se-pubescent; calyx lobes caducous; free central

column présent. Seeds ca. 0.9 mm long, comose

at both ends.—Fig. 10, UA-F.

JUVENlLt. FOI.IAGE. —Hopkins & Bradford 5038,

1 m high sapling: leaves 3-5 foliolate; leaflets

chartaceous with serrate margin; stems, leal

rachis and underside of the ntidrib pubescent;

stipules large, persistent, toothed. Small shrubs

in open, disturbed areas had leaves reladvely far

down the stems, the lower unes usually trifoliol-

ate and upper ones simple, sometintes with long

internodes (ca. 10 cm).

BreediNc, SYS TEM. —Polygamodioecious.

FlEl.D CHARACTERS. —Densely branched shrub

or small compact tree 1-71-20?) m high. Flowers

white, the calyx and ovary pale green or white;

petals and filaments white, stigmas tinged

purple; dise lobes dark red and strongly contrast-

ing with pale corolla; anther.s pink or cream.

Female inflorescences are comparatively small

and inconspicuous, scarcely projecting beyond

the foliage while male inflorescences are longer

and exserted beyond the leaves (j.C. B kaDTORD

pers. obs.).

Distribution and ecology. —Highland

régions of Viti Levu, Taveuni and Ûvalait; also

reported front Samoa. Recorded from dry open

forest, dense ridge scrub and windswept thickets

on mountain summit, escarpment edge and

swamp, at (360-)700-l4l0 m on Viti Levu, 840-

1100 m on Taveuni and 500-600 m on Ovalau.

Common on summit of Mt. Victoria

(Tomanivi). The form with large persistent stip¬

ules occurs in the Namosi District of Viti Levu

and on Ovalau.

In Match 1996 we found this species at three

localities, always on the highest peaks and ridges

in the vicinity and sometimes in boggy areas.

Near Monasavu Dam, Viti Levu, in disturbed

forest its associâtes included Pullea glabra and

Spiraeanthemuni serratum (Cunoniaceae), Rubus,

Vaccinium , Melastomataceae, Zingiberaceae,

Lycopodium , ttec feins and Gleicheniaceae (see

Hopkins & Bradford 5 021). On the summit of

Des Voeux Peak, Taveuni, stérile shrubs 1-3 m

high were one of the dominant components of

disturbed areas near the telecom tower (Hopkins

dr Bradford 5035 , 5039) with Scaevola

floribunda. It was also relattvely common in

undisturbed cloud forest with abundant

Freycinetia along the ridge towards Lake

Tagimaucia. Weinmannia affinis showed less syn-

chrony in its reproductive behaviour than W.

richii. Both within populations and with in indiv-

iduals we observed inflorescences at several stages

of development including matute flowers.

Wbere W. affnis occurs at the saine locality as

other Weinmannia species, there appears to be

sortie ccological séparation between them. For

instance, near Monasavu Dam, W. affinis was

found as a small tree or shrub in fiiirly short

forest near rhe road at its highest élévations

(1070-1210 m) and W. richii was seen as a 20 m

high tree in taller forest at lower élévation (ca.

840 m) in rhe same general area. On Des Voeux

Peak, Tavenui, W. affmis was again found onfy

near rhe summit and along tidges (ca. 1100 m)

as a small tree, shrub or epiphyre while mature

rrees of W. vitiensis were only seen below 900 m

in taller forest on the mountain slope.

Selkchd COLLECTIONS (from a total of 30 stud-

ied). —Fiji: Viti Levu: Bradford 591 , Tomanivi (Mt.

Victoria), trail from Navai to ridge crest below sura-

mii, I7°38’S-178 C 0I'E, fl.. 30 Mar. 1996 (MO, P,

SUVA); Gibbs 642 , Nadariv.tru. 2700 fi., fl., Sep.

1907 (BISH, BM, Kl; Hopkins & Bradford 5022, nr.

Monasavu Dam, road S from dam, 1210 m. fl., 29

Mar. 1996 (MO, P, SUVA); Koroiveibau DA 19184,

Ba, Mt. Evans Range, Natualevu, 3000 ft., fl., 8 Apr.

1965 (BISH, SUVA); Kmvivalniu & Qoro DA 14574,

Namosi, Mt Nadobc, 2S25 fr., fl., 16 Nov. 1965

(GH, L, SUVA); Koroiveibau DA 14 702, Namosi,

Mt. Vakarogosiu, summit 3400 fr., fr., 23 Nov. 1965

(K, SUVA); Kurutmli DA !3890, Nausori Highlands,

1900 ft.. fr.. 20 Julv 1964 (K, SUVA); karham DA

2201, Namosi. Korobasaliasaga, summit, 3950 ft., 11.,

24 Mar. 1940 (A. BISH, SUVA); Dirham DA 2598

pro parte, Namosi, Voma Peak, 2000 or 3000 ft., fl.,

23 Junc 1939 (A, BISH, SUVA); Smith 4905, Mba

ADANSONIA. Sér. 3 • 1998 • 20(1)

Hopkins H.C.F.

(formerlv Tholo North), nr. Nandarivatu, 800-

900 m, Fr'., 26 J une-19 Oct. 1947 (A, B1SH, K, L, P);

Thomerson DA 19527. Nandronga-Navosa, Nausori

Highlands, 850 m. H., 29 Oct. 1985 (SUVA);

Webster & Hildreth 14204, Mba. 1 uvua, summit Mr.

Victoria (Tomauivi). 4300 ft., fr., 13 July 1968 (GH,

SUVA). Taveuni: J. & U7 As h DA 19957 , Lake

Tagimauda, Somosomo, 850 m, fr., 15 Sep. 1982

(SUVA), Hopkins d r Bradjord 5036, Des Voeux Peak,

nr. telecom mtver and ridge towards crarer & lake,

1000-1100 m, buds. 2 Apr. 1996 (MO, P, SUVA);

Seematm 200, V'nn:i (fide Siniih), fr., 1860 (BM. GH,

K, P); Smith 8/8, borders ol Lake E of Somosomo,

700-900 m, fr,, 29 Dec. 1933-8 (an. 1934 (BlSH, K).

Ovalau: Smith 7608. summit Mc. Ndelaiovalau, 575-

626 m. fr., 1 1-25 May 1953 (BfSH, GH, K, L, P.

SUVA); Smith 7704 , su ni mil Ml. Tana I.ailai, 500-

550 m. fl., 1 June 1953 (BlSH, GH. L, P, SUVA).

LocAI. NAMES. —Katakata ( Smith 4905)', Vure

( Gillespie 2736).

Variation. —The size of the leaves and the size

and persistence of the stipules are variable but

the material cannot be readily divided into more

than one taxon (see SMITH 1985). The majority

of collections hâve small or medium sized, ±

coriaceous leaves and small, caducous stipules

(Fig. I IB). A few collections hâve larger leaves

and some of rhese hâve large, persistent stipules

that are rounded or pointed at the apex (Fig.

11A,C), e.g Smith 7 608 front Ovalau, Parham

DA 2598 pro parte and Korniveibau DA 14702

from Namosi région, Viti Levu.

4. Weinmannia vitiensis Seem.

Fl. Vit.: 110 (1866); Engl., Nat. Pflanzenfam., ed. 2,

18a: 256 (1930); A.C. Sm„ J. Arnold Arbor. 33: 135

(1952); Bernardi, Bot. Jahrb. Syst. 83: 207, t. 38

(1964); J. Parhant, PI. Fiji Islands: 125 (1972, rev.

ed.); A.C. Sm., Fl. Vit. Nova 3: 22 (1985).— Type:

Seemantt 199, Fiji, Kadavtl (holo-, K!, photo US!; iso-,

BM!, P!, also G, GH, MF.L, NSW jidc HOOGIANO).

Shrub or trec 3-20 m high. Branching often

dichotomous. Stems woody, somewhat thickened

at nodes, annular scar visible, internodes ca. 0.7-

3 cm long. Young stems minutely or sparsely

strigose, older stems glabrous with nutnerous

prominent lenticels, axillary buds densely velut-

inous, acute at apex. Stipules fugaceous even in

végétative material, ligulate to shortly spathnlate,

apex rounded to acute, to 3 mm long, minutely

hairy to ± glabrous. Leaves usually trifoliolate,

rarely simple, total length up to 8 cm including

petiole of 0.9-2 cm; petiole semîterete and nar-

rowly winged, especially distally, midrib promin¬

ent above, glabrous; latéral leaflets elliptical

(2,1-)3.3'4.7 X (0.8-)l-1.6 cm, unequal at base,

acute at apex; terminal leaflel elliptical or some-

times tiarrowly obovate, often larger and broader

chan the laterals, 4.3-6.5 X 1 3-2 cm, base atten-

uate, apex acute; leaflet blades glabrous, coria-

ccous, punccate below; margin sometimes

thickened, crcnate, the crenations acroscopic, 11-

13 on each sidc ofa leaflet; midrib slightly raised

above and prominent below, secondary and ter-

tiary venalion slightly raised on both surfaces,

often drying paler tlian intervenium.

Inflorescence a central rtiad or pentad witli the

lower racemes either in axils of leaves or not, or

in the axils of reduced leaves, sometimes also

with a pair of latéral triades; sometimes a few suc¬

cessive nodes on a shoot producing partial inflo¬

rescences simulraneously; latéral auxiFiary buds

sometimes présent at lower nodes of inflorescen¬

ce; pedunclé and rachis segments 0.7- 1.3 cm

long, sparsely pubescent, racemes 4-7 cm long.

Floral buds inserted singly; floral braers to l mm,

cymbiform or dagger-shaped, almost glabrous,

caducous. Flowers unisexual; pedicel 0.2-3 mm

long, almost glabrous; calyx lobes 0.6-0.8 mm

long, glabrous; petals 0.9-1.5 mm long; dise

lobes 0 3-0.8 mm long; in male flowers: fila¬

ments 2-2,5 mm long, the anthers 0.5 mm long

and ovary ça. 1 mm long, glabrous, the styles

0.2-0.3 mm long; in female flowers. the fila¬

ments to ca. 0.7 mm, ovary (post anthesis) 1.1-

1.5 mm long, sparsely strigose, the styles ca.

0.5 mm long.

Capsules 1.8-2.5 X 0.9 mm at dehiscence, the

styles 0.5 mm or less long, the exocarp sparsely

strigose, with minute longitudinal ridges, calyx

lobes not persistent, central column présent.

Seeds ca. 0.7 mm long, ca. 16 per capsule, como-

se ar both ends.—Fig. 10, 1 1G-K.

JUVENILE FOLIACE. —Differs markedly from

adult: see Hopkins & Bradford 5042, from a

sapling 3.5 m rail growing in shade, Taveuni:

ADANSONIA. sér. 3 • 1998 • 20(1)

Weinmannia in New Guinea and the western Pacific

Fig. 11.—A-F, Weinmannia affinis: A. shoot with large simple leaves and persistent stipules; infructescence a triad with reduced

leaves at node; B. shoot with small simple leaves, the stipules caducous; C, stipule trom A; D, male flower; E, temale flower; F,

dehisced capsule. (A, C, F, Smith 7606\ B, Seemann 200\ D, Gibbs 642 ; E, Parham 2201) - G-K, Weinmannia vitiensis : G, shoot

with an immature inflorescence (penlad) and reduced leaves at two nodes; H, leaf with slightly winged petiole; J, section of inflores¬

cence axis showing flower buds inserted singly; K, dehisced capsule. (G-K, Bryan 317). —Drawn by C. Monnié.

ADANSONIA. sér 3 • 1998 • 20(1)

Hopkins H.C.F.

leaves imparipinnate with 2-4 pairs of leaflets;

leafler margins crcnare; pétioles terete (not wing-

ed) with dense, erect hairs; stipules large, almosr

round, roothed.

BreHDING SYSTEM.—Dioecious. Flowers uni-

sexual but little material is available.

FlEl.D CHARACTKRS.—Dense shrub or tall,

canopy tree, sometimes gnarled. Bark rough,

scaly, ligbt grey; sapvvood orange-brown, coarse;

heartwood yellow-brown, moderately hard.

Flowers whire with a slight fragrance; pedicel and

calyx pale green, corolla and filaments white,

anthers cream, dise lobes dark brown-red

Distribution and ecology.—K nown only

front three of the smaller Fijian high islands at

300-900 m Habitats include tall, dense forest on

mountain slope (Hopkins & Bmdjbrd 5041), and

forest on sumntit ridge (Bryan 317); also in open

areas among bracken; loeally frequent.

MATRRIAI EXAMINl-.lt. —Fiji; Kadavu : Seemann 199,

buds, 1860 (BM. K, P), type. Taveuni; Hopkins &

Bradfon{ $041, Des Voeux Peak, W side. 900 ni, fl., 2

Apr. 19% (MO, P, SUVA); Smith 8401, hilb F of

Somosomo, W of old crater, 660-900 m, su, 18 Aug.

1953 (BIS H, GH, K, JL P, SUVA). Mon ht Bryan

317, summit ridge, 300 m, buds, If., 11 Joly 1924

(BISH, K); .Smith 1334, Ndelaimoala, fr , 20-24 Mar.

1934 (BISH, K, P); Harvey s.n., Feegee (sic) Islands,

s.loc., Nov. 1855 (K).

Local n.aml.— Molau ndamu (Smith 1354).

RelatIONSHIPS.—R egarded by SMITH (1985)

as not sharply distinct front Weinnuinnia richii ,

but the structure of the racemes and the infloresc¬

ences are quite different and rhese species belong

to different sections of the genus. On présent

evidence there is no overlap in the distributions

of W. vitiensis and IL. richii.

Besides the différences in the leaves and habitat

between Weinnuinnia vitiensis and IL) afjinis, in

the former the apical buds of the shoots abort

less offert so tltat dichotomous branching is less

frequent.

BernakDI (1964) included Weinnuinnia raia-

teensis J.W. Moore (front Raiatea, Society

Islands) in IL', vitiensis. Both hâve trifoliolate

leaves with a narrowly winged rachis; the leaves

of W. raiateensis, however, tend to be smaller and

less coriaceous than those of W. vitiensis.

Doubtful species

Weinmannia spiraeoides A. Gray

U,S. Expi. Exped., Phan, 1: 677 (1854); Seem., Fl.

Vit.: 110 ( 1 86 S ). Engl., l.innaea 36: 644 (1870), Nat.

Pflanzenfam., ed. 2, I8a: 256 (1930): A.C, Sm., J.

Arnold Arbor. 33: 133 (1952); Bernardi, Bot. Jahrb.

Sysi. 83: 208 (1964); J. Parbam. PI. Fiji Islands: 125

(1972, rev. ed.); A.C. Sm., Fl, Vit. Nov. 3: 20

(1985).—Type: US 48073, l iji (Feejee) Islands,

Ovolau, U.S. Expi. Exped. under the comniand of

Capt. Wilkes (liolo-, US 480731).

Known only front the stérile type whiclt has

chartaceous, 3-5-foliolate leaves with dentate

margins and toothed stipules. It appears to be

juvénile foliage of Weinmannia as suggested by

SMITH (1985). However, immature foliage is so

variable as to preclude identification in rnost

cases (see I foPKINS 1998a) It is unlikely that this

name représente a distinct taxon.

V. WEINMANNIA IN SAMOA AND THE

COOK ISLANDS

Weinmannia has been collectcd front Savai’i

and 'Upolu in Western Samoa and front Tutuila

and the Manua group (Ta'u and Olosega) in

American Samoa. It also occurs on Rarotonga in

the Cook Islands to the SE but is not known

from Tonga or Niue. Three species hâve been

describcd font this région: W. samoensis A. Gray,

W. manuana Christoph., and W. rarotongensis

Hemsl. in Chceseman (Hopkins & Florence

1998, Fig. 7F-M); ail bclonging to sect.

Leiuspermum. Weinmannia ,/[finis -and W. richii

from Fiji hâve also been reported

(Chrimophlrsln 19.38; Bf.rnardi 1964).

Three treatments of Weinmannia hâve been

published for this région, none of them satisfact-

ory. CHRISTOPHKRSF.N (1938) recognised four

species: VF 7 , manuana, W. ajjinis, W. richii and

W. samoensis and did not discuss W.

rarotongensis. SMITH (1952) constdered ail the

species in Samoa to be endemic and recognised

W. samoensis, W. manuana and Weinmannia sp.

(based on Chnstophersen 787, 2561 , 2735, ail

stérile); Christophersen 534 (also stérile) was

100

ADANSONIA. sêr. 3 - 1998 - 20(1)

Weinmannia in New Guinea and the western Pacific

unplaced; W. rarotongensis was separated from

W. samoensis by implication. BERNARDI (1964)

placed W. manuana into synonomy with W.

affinis , included W. rarotongensis in W. samoensis,

and recognised W. richii, based on Christophersen

534, 787 and Rechinger 1647, Thtts none of his

species was endemic ro Samoa.

A. WHlSTLtR (pcrs, comm.) and J.C,

Bradford (pers. comm.) hâve seen Weinmannia

growing in Samoa, -and both çonsidér rbat there

is only a single, variable species in this group of

islands, whose leavcs can therefore be unifoliola-

te, trifoliolate or imparipinnate. BRADfORD

(pers. comm.) found that at lower élévations,

trccs usually had 3-7 leaflets per leaf and ar

higher élévations, 1-3 leaflets per leaf.

From srudies of herbarium spécimens, I hâve

failed to reach a conclusion about the taxonomy

of species from Samoa and the Cook Islands, and

to avoid making mattcrs worse, will comment

only on ihe possible occurrence of the Fijian spe¬

cies. Ail the trifoliolate collections 1 hâve seen

from Samoa referred by BliRNARPI and

CHRISTüri lliRSHN to Weinmannia richii are stérile

and likely to be juvénile specimens of other taxa.

1 hâve seen no fertile material from Samoa which

could be equated with W. richii , which should

therefore be considered endemic to Fiji. Several

specimens with small. unifoliolate leaves from

Samoa cannot be separated from W affinis

although many of the unifoliolate collections fall

outside the range of the Fijian material. It is thus

not clear at présent whether W. affinis is endemic

to Fiji or not.

The structure of the inflorescence of ail collect¬

ions seen from Samoa and Rarotonga is typical

for sect. Leiospermum.

VI. WEINMANNIA IN NEW CALEDONIA

(with R.D. Hoogland')

Four endemic species of Weinmannia occuc in

New Caledonia (Hoogland, unpublished

manuscript at P). Full synonomy, descriptions

and citation of specimens will be published in

the “Flore de la Nouvelle-Calédonie” and only a

summary is given here. The taxonomie changes

indicated here should be attributed to

HOOGLAND, and in the citation of types below, !

shows that a specimen lias been seen by him.

Ail the species in New Caledonia belong to

sect. Leiospermum and appear more closely relat-

ed to one another than to other members of the

section. They hâve small, caducous, usually ligul-

ate stipules in mature fohage; juvénile f’oliage

may hâve more persistent, salvc-rform stipules.

The flowers are bisextial and white, and the calyx

lobes usually fall in fruit. In Weinmannia dichoto-

ma, W. paitensis and W. ouaiemensis , végétative

growth often involves dichotomous branchmg,

and in the inflorescence, the apical bud is often

aborted. When this is not so, the apical and axill-

ary buds usually develop equally. Latéral auxiliary

buds are especially prominent in the infloresc¬

ences of W, dichotoma (see Hopkins 1998a,

Fig. 3M).

Key to the species of New Caledonia

1. Leavcs on reproductive branches alwavs simple ........2. W. paitensis

T. Leaves on reproductive branches trifoliolate ...2

2. Végétative growth usuallv by development of both apical and axillary buds, i.e. branching not dichoto¬

mous; inflorescence usually a central, apical triad (or sometimes a pentad), often with two latéral triads ....

... 1. W. serrata

2’. Végétative growih almosc exclusive!/ by development of shoots from latéral, axîllary buds, the apical bud

aborted, i.e. branching diçhotomou», though the branches sometimes unequal; inflorescence usuallv a pair

of latéral dyads or triads, the apical bud aborted (tarcly otherwise).........3

3. YoUng shoots sparsely hairv to glabrcsccnt; upper surface ol leaflets glabrous; pedicel 0.3-0.5 mm diam-

eter, glabrous or bricrly and sparsely hirsute; filaments 2 5-4.5 mm long; ovary sparsely strigosewith 8-12

(-14) ovules per locale; styles 2-3 mm long ..... 3. W. dichotoma

3’. Yoting shoots and inflorescence axes densely tomentose; upper surface of leaflets loosely histuic, glabrcsc-

ent; pedicel 0.5-0.7 mm diameter, moderately hirsute; filaments 5.5-6 mm long; ovary strigose-hirsute

with 12-16 ovules per locule; styles 2-3 mm.4. W. ouaiemensis

ADANSONIA, sér. 3 • 1998 • 20(1)

101

Hopkins H.C.F.

1. Weinmannia verrata Brongn. & Gris

Bull. Soc. Bor. France 9: 73 (1862).—Type:

Vieillard 572, Hab. in montibus Novae Caledoniae,

prope Balade (holo-, PI; iso-, NSW!).

Végétative branching often dichotomous.

Leaves subcoriaceous, not punctate below.

Inflorescence usually a central triad or pentad,

often with latéral triads devcloping from axillary

buds (i.e. apical bud reproductive, rarely abort-

ed); reduced leaves sometimes présent at nodes

within the inflorescence.

Tree up to 25 ni, widesptead on Grande Terre

in humid forest and gallery forest between 150-

800 m, on greywaekes, schist and basait.—Fig.

12A-E.

2. Weinmannia paitensis Schltr.

Bot. Jahrb. Syst. 39: 124 (1906).—Type: Schlechter

14941 , New Caledonia, Sud-Bezirk: auf dern Gipfêl

des Mont Mou bui Paita, ta 1230 ni (holo-, B; iso-,

BM, BR, E, G, K, U, LUI. LE, M, NSW!, P!, W, Z!

yw&BitkNAli£M and Hoogi and).

Weinmannia fthornei Guillaumin in Guillaumin et al.,

Stud. Nat. Hîsr. lowa Univ. 20: 31 (1965).—Type:

Thorne 28733, New Caledonia, Mt. Mou, 1050 m,

8 Nov. 1959 (holo-, PI; iso-, L!, P!, RSA, Z); syn.

nov. of HüOGLAND.

Végétative branching usually dichotomous;

dichotomous branching is fixed in juvénile plants

while in adult foliage, branching can be dichoto¬

mous or not (J.C. BRADFORD pers. comm.).

Leaves on reproductive shoots simple but foliage

on lower branches within the crown trifoliolate.

Leaves coriaceous, often punctate below.

Inflorescence a pair o! latéral dyads or triads, the

apical bud aborted; dyads without buds between

the racemes.—Fig. 12F-H.

A small tree known from Mt. l'Aoupiné and

several mountains towards the south, including

Mt. To, Mt. Kouakoud, Mt. Ouin and Mt. Mou,

in dense humid forest from (500-)950-1300 m,

on ultrabasic or schistose rocks.

3. Weinmannia dichotoma Brongn. & Gris

Bull. Soc. Bot. France 9: 73 (1862).—Type:

Vieillard 569, Hab. in montibus Novae Caledoniae,

prope Balade (lecto- of Bl-ltNARDI, P!: isoiecto-, P!).

Weinmannia monticnla Daniker, Vierteljahrsschr.

Naturf. Ges. Zurich 76 = Mitt. Bot. Mus. Zurich

137: 165 (1931).— Type: Diiniker 1812, New

Caledonia, auf den flanchen Hohenrucken an der

Westflanke des Ignambi. 6 June 1925 (holo-, Z!);

syn. nov. of Hoogland.

Végétative branching largely dichotomous.

Leaves on reproductive shoots trifoliolate, those

on lower leaves and immature foliage 3-7-fotiola-

te. Leaves coriaceous, often punctate below.

Inflorescence a pair of dyads or sometimes triads

in larger-leafleted specimens, the apical bud

aborted. Auxiliary buds at base of peduncle, in a

plane perpendicular to that of the leaf-bases,

large and conicak

Shrub or small tree up to 12 m. Widespread on

Grande Terre in humid forest on mica-schist at

400-1550 m and ultrabasic substrate between

950 and 1300 m.

BERNA RDI (1964) regard ed W. monticnla, with

larger leaflets, as distinct from W, dichotoma but

there are numéraux intermédiares. The possibil-

ity of any ecologicat différences correlated with

leaflet si/e nceds to be investigated in the ficld.

On Mt. Mou, the shape of the leaves

approaches rhat of W. semitil , wirh a Lilly sinu-

ate margin. On both Mt. Ouin and Mt. Mou it

grows in close proximity to W. paitensis in

Nolbofagus cloud forest on deep, peaty soil

(Bradford &c Hopkins pers. obs.). The imma¬

ture foliage of these two species can be diffîcult

to separate aLthough mature canopy foliage is

clearly distinct.

4. Weinmannia ouaiemensis

(Guillaumin & Virot) Lloogland, comb. nov.

Cunvnin ouaiemensis Guillaumin & Virot, Mém.

Mus. Natl. Hist. Nat., sdr B, Bot. 4: 28 (1953).—

Type: Virot 751, New Caledonia, Arête rocheuse

menant au Mt. Mi (versant W) point culminant du

massif de la Roche Ouaième, 1050 m (holo-, P!; iso-,

A, P! 2 sheets),

Végétative branching dichotomous. Leaves tri¬

foliolate, the petiole not winged; blade coriac¬

eous, punctate below, the margin revolure; white

102

ADANSONIA, sér, 3 • 1998 * 20(1)

Weinmannia in New Guinea and the western Pacific

deposit on upper surface of old leaves.

Inflorescence a pair of latéral dyads (without a

bud between the racemes) or triads, the apical

bud usually aborted or rarely developed into a

dyad.

Stunted shrubs endemic to Roche Ouaième, in

exposed thicket and low humid forest on mica-

schist, between 700-1150 ni.

VII. WEINMANNIA IN NEW ZEALAND

There are two closely related allopatric species

occurring in New Zealand: Weinmannia racemo-

sa L. f. occurs on Sourh and North Islands, as far

north as Hamilton, and W. sylvicola Sol. ex A.

Cunn. is found on North Island, to the north of

Hamilton (BF.RNARDI 1964; WARDI.F. 1966).

These two .species appear to lie closely related

and are superficially similar to one another, but

the adult leaves of W. racemosa are usually simple

(Fig. 13G) while chose of W. sylvicola are usually

trifoliolate (Fig. 13A,C) (and sec Wardlf. for

further différences).

In borh species the Flowers are inserced singly in

the axils of btacts on the racemes, as in other spe¬

cies of sect. Leiosùerrnum Other characters char

they share with most species of this section are;

leaves often punctate below; leaf rachis slightly

expanded; stipules caducous and wliere présent,

ligulate not orbicular (Fig. 13L); calvx lobes fal-

len in fruit (Fig. 13F,J) (a few exceptions); calyx

lobes glabrous or sparscly stngose. These species

aiso show a number of features which are not

widespread within the section. The Flowers are

somewhat perigynous (Fig. 13E) (rather than

hypogynous), and they may hâve a greater tend-

ency to be 5-merous or to hâve a 3-merous ovary

than other Pacific species. The Flowers are

bisexual (Fig. 13D.FI), as in the New Caledonian

species, where as in most Pacific species the

majority of inflorescences hâve unisexual flowers.

The inflorescences differ somewhat from those of

other Pacific species and are described by

Hopkins & Bradfoiui (Hopkins 1998a).

Acknowledgements

The work by HCFH was financed by the Human

Capital and Mobility Programme of the European

Union via the network on Botanical Diversity of the

Indo-Pacific. J,C. BkadforD is grateful for Financial

support from Washington University, including the

NSF Graduate Research Training Support Grant NSF

B1R-9256779, and from the Missouri Botanical

Garden, especially through the Mellon Foundation

Fellowship. In connection with our fleldwork in

1996, we are greatly indebted to M. DOYIF, (SUVA,

Fiji) and the Fiji F.lcctricirv Atithority (Monasavu

Dam) the Dircctnrs ot the “Service de l'Environ¬

nement" for the North and South Provinces in New

Caledonia, B. Suprin, and T. JAPPRC. and his sraff at

ORSTOM in Nouméa for their lieip. We rhank the

duectors of the herbaria cited fbr access to spécimens,

and C. MoNNif and J. Saussoi i i -GuErm for the

illustrations, HCFH rltanks M. Coodf for corrccting

the Latin diagnosis for Weinmannia croftii. M. LOCK

and P, I.OWRY for helpful comments on the manus-

cript and E. ARMÈDE for help with typing.

REFERENCES

BERNARDI I.. 1964.—Rcvisio generis Weinmanniae.

Pars III: Sectiones 111 - IV - V -VI. Bol. lahrh. Syst.

83: 126-184; 185-221.

QIKISTOPHERSEN E. 1938.— Flawerinvplants of Samoa

- II. Bernice P. Bish. Mus. Bull. 154)

David A. 1995.— The Voyage of H MS Heriild.

Meigunyah Press of Melbourne University, Carlton,

Victoria.

HOPKINS H.C.F, 1998a.—A révision of Weinmannia

(Cunoniaceae) in Malesia and the Pacific. 1.

Introduction and an account of the species of

Western Malesia, the fesser Sunda Islands and the

Moluccas. Adansonia, sér. 3, 20: 5-41.

Hopkins H.C.F. 1998b.—A révision of Weinmannia

(Cunoniaceae) in Malesia and the Pacific. 2. Sulawesi

and the Philippines. Adansonia , sér. 3, 20: 43-66.

Hopkins H.C.F, A FiokKnce J. 1998.—A révision

of Weinmannia (Cunoniaceae) in Malesia and the

Pacific. 4. The Society, Marquesas and Austral

Islands. Adansonia. sér. 3, 20: 107-130.

l’ARHAM H.B.R. 1941.—Fiji Plants, their names and

uses .J.'Polyncs. Soc. Suppl. 16: 65.

PERRY L.M. 1949.—Plantae Papuanae Archboldianae,

XIX. ,/ Arnold Arbot 30: 139-165.

SCHLECH l ER R. 1914.— Die Cunoniaceae

Papuasiens. Bot. fahrb, Syst. 52: 139-166.

Sc I n rc htp.r R. 1917.—Cunoniaceae, Nova Guinea

12: 491 -493.

SMITH A.C. 1952.—Studies in Pacific Island Plants,

XII. The Cunoniaceae of Fiji and Samoa. J. Arnold

Arhor. 33: 119-149.

Smith A.C. 1985.—Cunoniaceae. Flora Vitiensis

Nova 3. Pacific Tropical Botanical Garden, Lawai,

Hawaii.

STAFLEU F.A. & Cowan R.S. 1985.— Taxonomie

ADANSONIA, sér, 3 • 1998 • 20(1)

105

Hopkins H.C.F.

Literature, ed. 2, vol. V: Sal-Ste. Regnum vegetabile,

vol. 112.

Turrill W.B. 1915.—A contribution to the flora of

Fiji. / Linn. Soc. Bot. 43: 15-39.

WARDLE P. 1966.—Biological Flora of New Zealand

1. Weinmannia racemosa Linn. f. (Cunoniaceae).

Kahahi. New Zealand J. Bot. 4: 114-131.

WHEATLEY J.I. 1992.— -A guide to the common trees of

Vamiatu. Dept. of Forestry, Republic of Vanuatu.

WHITMORE T.C. 1966.— Guide to the forests of the

British Solomon Islands. Oxford Üniv. Press,

London.

Manuscript received 30 October 1997;

revised version accepted 27 February 1998.

106

ADANSONIA, sér. 3 • 1998 • 20(1)

A révision of Weinmannia (Cunoniaceae)

in Malesia and the Pacific. 4. The Society,

Marquesas and Austral Islands

Helen C.F. HOPKINS

Laboratoire de Phanérogamie, Muséum national d'Histoire naturelle,

16 rue Buffon, 75005 Paris, France.

Masons Arms, Hutton Roof, via Carnforth, Lancashire, LA6 2PE. U.K.

Jacques FLORENCE

ORSTOM, Laboratoire de Phanérogamie, Muséum national d'Histoire naturelle,

16 rue Buffon, 75005 Paris, France.

florence@mnhn.fr

KEYWORDS

Weinmannia,

Cunoniaceae,

Society Islands,

Marquesas Islands,

Rapa Island.

ABSTRACT

Six species are recognized in this région, ail belonging to sect. Leiospermum.

Weinmannia rapemis is confined to Rapa in the Austral Islands and its report-

ed occurrence in Pitcairn is discussed. Weinmannia panùflora occurs on niost

of the large islands of the Societies except Raiatea and its leaf shape and

indumentum vary with its ecology. Weinmannia vescoi and W. miateensis are

both confined to Raiatea. In the Marquesas, W. manjnesana is rcprcsented by

two varieties and a distinctive, new species, W. tremuloides, is described and

illustrated front Faut Hiva.

MOTS CLES

Weinmannia,

Cunoniaceae,

Iles de la Société,

Iles Marquises,

Rapa.

RÉSUMÉ

Six espèces, appartenant à la sect. Leiospermum, sont reconnues pour la

région. Weinmannia rapensis est confiné à File de Rapa dans les Australes et

sa présence à Pitcairn est discutée. Weinmannia parviflora se rencontre dans

la plupart des grandes îles de la Société, sauf à Raiatea, avec une grande varia¬

bilité dans la forme des feuilles et la pubescence. Weinmannia vescoi et W.

rauileensis sont tous deux propres à Raiatea. Pour les Marquises, W. manjue-

sana compte deux variétés et une espèce nouvelle, W. tremuloides, endémique

de Fatu Hiva, décrite ici et illustrée.

ADANSONIA, sér. 3 • 1998 • 20(1)

107

Hopkins H.C.F. & Florence J.

In SE Polynesia, to tlie east of the Cook Islands,

Weinrnannia is fourni in the Society Islands, the

Marquesas Islands, and on Rapa in the Austral

Islands. The genus is a significant component of

the végétation on Tahiti and on several of the

Marquesas, in contrast to much of the Malesian-

Paciflc région where it is generally focally com-

mon at most.

With the exception of Weinrnannia rapensis

which is morphologically distinct and geographic-

ally isolated, material from this région of the

Pacific is rather variable, especially in the shape,

size and texture oi the leaves. This treatment

recognises five species from the Society and

Marquesas Islands, including a new one from

Fatu Hiva (Table 1).

The leaves of the newly named species are very

distinctive but the remaining taxa are inclined to

grade into one anothcr to somc extent, and the

characters that difterentiate between them often

overlap or are nor completely diagnostic.

Provenance must therefore be used occasionally

in identification. While this is not generally a

recommended taxonomie practice, the under-

lying morphological and geographical patterns

would otherwise be lost

For example, Weinrnannia parviflora appears to

be confinée! to the Society Islands but a few spé¬

cimens from the Marquesas (belonging W mar¬

quesana var. marquesana) are vegetatively difficult

to distinguish from it. Most collections of W.

raiateensis hâve small, délicate leaves but a few

hâve larger ones and/or some unifoliolare leaves,

and rhus resemble some specimens of W. parvi¬

flora from Tahiti. Material front l luahine, desi-

gnared here as W. parviflora , is somewhat

interrnediate between that species and W. raia¬

teensis (in having caducous stipules and winged

pétioles when the leaves are trifoliolare).

A large number of recenr collections, especially

by J. FiüRFNCT, provide a derailed picture of

variation in relation to ecology and distribution.

Weinrnannia parviflora and W. marquesana espe¬

cially show phenotvpic variation correlated with

altitude and exposure. At mid élévation on Tahiti,

W parviflora has narrowly elliptical leaves with a

crenate-sinuate rnargin, while at higher altitude

the leaves are shorter and often ovate, the stems

Table 1.—The species of Weinrnannia in the SE Polynesia according to recent treatments.

Current révision

Bernardi (1964)

Fosberg & Sachet (1972)

Florence (1982)

Society Islands

W. parviflora

W. parviflora var. parviflora

(by implication)

W. parviflora

var. parviflora

W. vescoi

(syn. W. ovalifolia)

W. vescoi

(syn. W. ovalifolia)

W. vescoi

W. raiateensis

in synonomy under

W. vitiensis

W. raiateensis

Marquesas Islands

W. marquesana var. marquesana

W. marquesana

W. parviflora var. marquesana

+ W. parviflora var. glabrata

W. marquesana var. myrsinites

W. parviflora var. myrsinites

W. tremuloides

Austral Islands

W. rapensis

108

ADANSONIA. sér, 3 • 1998 • 20(1)

Weinmannia in the central Pacific

are thicker and ofren more densely pubescent, the

internodes are shorrer and the leaf margin is

alrnost entire. A similar pattern is seen, though

rarely, in W marquesana. lt appears that these

high altitude forms bave evolved in parailel on

Tahiti and in the Marquesas. Weinmannia parvi-

flora is absenr Irom Raiatea and material with

rounded leaves and short, glabrous internodes ( W.

vescoi) is nor obvîously derived ftom any other

taxon. Thus on Raiatea, Tahiti and some of the

Marquesas, there are specimens Irom reladvely

exposed, windswept sites ar high altitude with

ovate or rounded, coriaceous leaves and short

internodes, but rhey ail belong to different taxa

depending on the island on which they occur.

Weinmannia has small, comose, anaemochorous

seeds that appear to be well adapted to wind dis¬

persai. It is thus not surprising that représenta¬

tives of the genus are found on ail rhe high

volcanic islands of rhis région. The seeds ot W.

marquesana , W. raiateensis and W. vescoi are simil¬

ar to one another and unitsual in sect.

Leiospermurn in that the comose hairs are rela-

tively short, and shorter than those of W parvi¬

fiora from Tahiti. If the presence of these hairs is

related to dispersai, this could indicate a décliné

in the dispersabiliry of these three taxa. The Iles

sous le Vent (including Raiatea, Tahaa, Huahtne,

Bora Bora) and the Marquesas are older than the

Iles du Vent (Tahiti and Moorea), Tahiti being

about one million yvars old (see BROUSSE 1993).

A generic description applicable to the Pacific

species of Weinmannia, and general notes on

tnorphology, are givett in HOPKINS (1998a). Ail

the species of the région dealt with hcrc belong

to sect. Leiospermurn , whose characters are also

discussed in HOPKINS (1998a). An index to ail

the taxa of Malesian-Pacific Weinmannia (see

HOPKINS 1998a,b, c) is given at the end of this

paper.

Key to the species and varieties of Weinmannia in the C Pacific

1. Leaves usually trifoliolate ..2

1 ’. Leaves usually simple (unifoliolate) .4

2. Yottng stems * glabrous, pétiole usually glabrous; stipules nut recurved and usually caducuus .3

2’. Young stems densely pubescent, pétiole semiterete but not winged, pubescent; stipules strongly recurved

and usually persistent (Rapa) .........6, W. rapensis

3. Pétiole latfrally flattened and U-shapcd in cross secrion (Fatu Hiva) ...... 5. W. tremuloides

3’. Pétiole t fiat on adaxial surface, not latérally flarrened (Raiatea) ...2. W. raiateensis

(and rarely sonte specimens of W. parvifiora but not on Raiatea)

4. Young stents completel) glabrous (Raiatea) ..3- W. vescoi

4’. Young stems witlt sparse to dense indumentum ...5

5. Stipules usually persistent and elliptical, recurved, the apex rounded; capsules small, 2.5-3-5 mm long at

dehiseertcc (Society Islands) . I. W. parvifiora

5’. Stipules usually caducous, when présent usually ± round, conduplicate, rhe apex acute and the margin

sometimes toothed, or sometimes stipules elliptical-ligulare; capsules larger. 3.3-5 mm at dehîscencc

(Marquesas) ...6

6. Leaves elliptical or ovate, large (2.5-)3.3-7.5 X (l-)1.3-3.6(-4.4) cm, venation usually visible on adaxial

surface and réticulum on abaxial surface not dense .4a. W. marquesana var. marquesana

6’. Leaves ovate, smaller, 1.6-2.7(-3..3) X 0.7-1.7 cm, venation obscure on adaxial surface and réticulum on

abaxial surface dense...4b. W. marquesana var. myrsinites

1. Weinmannia parvifiora G. Forsl.

Fl. Ins. Austr.: 29 (1786); Willd., Sp. PL, ed. 4, 2:

438 (1799); Poir., Fncycl. 7: 580 (1806); A. Gray,

U.S. Expi. Exped., Phan. 1: 673, Atlas t. 85A, Fig. 1-

5 (1856) p.p.; Dccne., Voy, Vertus, Bot. Texte; 26,

Atlas t. 20 (1846); Seem., Fl. Vit.: 109 (1865); Engl.,

Linnaea 36: 647 (1870) pro parte, Nat. Pflanzenfam.,

ed, 2, 18a; 256 (1930); Bertiardi. Bot. (ahrb. Syst, 83:

195 t. 39 (1964); Fosberg & Sachet, Micronesica 8:

44 (1972).— Leiospermurn parviflorum (G. l'otsc.) D.

Don, Ediuburgh New Philos. J. 9; 91 (1830) pro

parte excl. spec New Zealand.— Merretia ntnninalis

Banks & Sol., mss. Murattia rernnnalis Sol, in

Parkinson, Drawtngs gf Tahit. PL, t. 98, ex Seem., FL

Vit.: 109 (1865), pro syn. sub Weinmannia parvifiora-,

Britten, J. Bot. 45: 315 (1907).— Weinmannia parvi-

ADANSONIA, sèr, 3 • 1998 • 20(1)

109

Hopkins H.C.F. & Florence J.

flora var. parvijlora-, implied by Fosberg & Sachet,

Micronesica 8: 44 (1972).—Type: Forster s.n., s.loc.,

Pallas Hb. (lecto- Itère designated, BM! pro parte excl.

left hand part of sheer).

Shruli or tree 1-10(-12) m high and up to

45 cm dbh. Branching usually not dichotomous

at lowcr élévation, more often dichotomous at

high élévation. Young stems usually densely

covered in erect, stiff hairs ca. 0.4 mm long,

sometimes glabrate; ai lowcr altitudes, internodes

up to 5 cm long; at high altitude internodes

usually short, 0.5-1.5 mm long, the stems often

thiclc, up to 0.3 mm diameter and nodes thicken-

ed, older stems glabrescent, ca 0.5 mm diameter

with numerous pale lenricels. Stipules usually

persistent at several nodes on reproductive shoots

(except Moorea and Huahinc), elhptical, spathul-

ate or ± orbicular, the margins recurved and the

whole stipule recurved, 0.4-0.5(-0.9) X 0.5-0.6

(-0.8) cm, the apex rounded, puberulem on both

surfaces, venanon usually indented above. Leaves

usually unifoliolate, sometimes mixed with trifol-

iolate ones. Unifoliolate leaves somewhat variable

in shape and texture according to alritude: at mid

altitude, subsessile to petiolate, the petiole ± tere-

te to stmiterete, 0,2-1 cm long, the blade usually

narrowly elliptical, 3.5-8,8 X 1.2-3(-3.5) cm, the

base either acute and decurrent into the petiole

or more rruncate with a sharp disrinction bet-

ween petiole and blade, apex acute, the blade

chartaceous, sometimes puncrate below, the mar-

gin sometimes thickened, crenate, dentate or

sinuate and sometimes undulate, 10-15 notches

on each side of a leaf; at higher altitude, shortly

petiolate, the petiole ca. 0.3 mm or 0.5 mm if

blade cuneate at base, the blade 3.1 -5-7(-7) x

( 1.5-) 1.9 X 3.3(-4.2) cm, ovate to oblong, the

base usually truncate, the apex broadly acute to

obtuse (rarely rounded), coriaccous or subcoriac-

eous, margin crenulate. Triloiiolate leaves up to

9 cm in length, the petiole 1-1.5 cm long; on

Tahiti the petiole terete to semiterete, the upper

surface not winged; on Moorea and Huahine,

the petiole narrowly winged; latéral leaflets acute

and unequal at base; terminal leaflet long-

attenuate at base.

Inflorescence a pentad (especially at high éléva¬

tion) or heptad, often with additional racemes

arising at the lower most node of the inflorescen¬

ce from latéral auxiliary buds; reduced leaves

sometimes présent at nodes within inflorescence;

peduttcles and rachis segments 0.5-4 cm long,

shorter at higbcr altitude, puberulem; racemes

up to 13-5 cm long at lower altitude and to 6

(-9.5) cm long at high altitude. Stipules at nodes

within inflorescence smailer than at végétative

nodes. Floral buds inserted singly; floral bracts to

0.4-0.7 mm long, subcarinate, strigose-pnberul-

ent, usually cadueous. Flowers usually unisexual,

rarely bisexual or ?protandrous: pedicel 0.8-

2.5 mm long, puberulem; calyx lobes 0.4-

0.9 mm long, puberulenr on ourer surlace;

corail a oblong to ± bluntly triangular, 0.7-1.6 X

0.5-1.5 mm; dise lobes 0.3-0.7 mm long; in

male flowers: filaments 1.9-3 mm long, the ovary

0.5-0.9 mm long, puberulem and rite style 0.1-

0.2 mm long, curved inwards; in female flowers:

filaments 0.7-1-1 mm long, die ovary > 1.1 mm

long ar antbesis, puberulenr, and tbc styles >

0.7 mm long, ± straight, the stigmas capirate and

papiilose.

Capïules 2.5-3.5 X 1.5 mm at dehiscence (up

to 0.5 mm longer prior to dehiscence), the styles

0.5-0.8(-l) mm long; the exocarp sparscly pub-

erulent to almost glabrous, endocarp sometimes

separating from exocarp in old fruits; calyx lobes

usually cadueous; central column présent. Seeds

0.8-1 mm long, comose at each end, the hairs

0.5-0.7 mm long.—Fig. 1A-K, 2.

JUVENILE FOLIACE (high altitude).—The seed-

ling anached to van Balgooy 1796 has simple,

chartaceous leaves, up to 6.5 X 5.5 cm, with the

margin dentate, the stipules tnuch as in the adult

fbliage and the stem ca. 3 mm thick, woody but

only sparsely hairy.

BrekdING SYS i KM. —Polygamodioecious.

While the flowers on most specimens are either

male or female, a few collections hâve racemes of

mostly male flowers with a few bisexual ones

mixed in (e.g. lloogLwd & Florence 12920), and

others hâve male flowers and fruit on the same

specitnen (Gagné & Montgomery 2362)

FlElD CHARACTERS. — At lower élévations, fre-

quenrly a small tree and ar high élévation, a

shrub or gnarled tree, the branches sometimes

clothed in bryophytes. Wood red, hard (Florence

& Varney 11093) or blaze pink and wood creamy

110

ADANSONIA, sér. 3 • 199B • 20(1)

Weinmannia in the central Pacific

Fig. 1.—A-K, Weinmannia parvifiora: A, flowering shoot, the leaves typical (or mid altitude ort Tahiti and the inflorescence a heptad

with additional racemes at the lowest node, developîng Iront latéral auxiliary buds; B. leaf, typical of plants from exposed

locations at high altitude on Tahiti; C. leaf, trom Moorea: D. trlfoliolate leaf, Tahiti; E, stipule with recurved margins; F, detail of

node with 2 opposite petloles and the peduncles of 2 partial inflorescences, the apical bud aborted; each peduncle with a

“collar" (remnants of fused stipules that protect the intlorescence in bud) and latéral auxiliary buds at base; G, maie flower;

H, female llower; J, dehisced capsule with persistent central column and one seed remaining in locule; K, seed. (A, E, G,

Florence 9090, B, St. John & Fosberg 17146; C, Florence 8365 ; D, H-K, Florence 9089, F, Florence 3110). —Drawn by C. Monnié.

ADANSONIA, sér. 3 • 1998 • 20(1)

111

Hopkins H.C.F. &c Florence J.

(van Balgooy 1685). I.caves pale green or bright

green above, shiny, toriaceous. Inflorescence axes

and calyx show a colour polymorphism at lower

altitudes, being eithei pale green or purple-red,

but above 1500 m only the red form was observ-

ed (J.C. BradfoRU pers. comm.). Buds white or

red. Flowers with pedicel and sepals pinkisb, pale

purple or green, petals and stamens white, dise

segments pale purple or dark, anthers yellow,

ovary white or pale green, die stigmas pinkish in

female Flowers (spécimen ficld labels and J.C.

BRADFORD pers. comm.). Reportcd as odourless

(Quayle 39) or sweetly scented and visired by

bees (J.C. BUADIORI) pers. comm.). Fruit green

sometimes ringed with red, reddish, or pale

purple; latcr brown.

Distribution and kcology. —Abundant on

Tahiti and also occurting on Moorea, Huahine

and Bora Bora [fuie FLORENCE (1982), who

reports tkat Dumont d'Urville 200 (s.l., s.d.) is

from that island). Also grows on Tahaa (J.

FlORENCL pers. obs.)<

On Huahine, confined ro rhe central crest bef-

ween Mt. Matoereere and Mt. Turi, from 450 m

to rhe summit (670 m), in forest dominated by

Metrosideros and Macaranga, growing with

Alstonia and Myrsine, and cspecially recorded

from ridges. On Moorea, flrst âppearing around

400-500 m, and co-dominant with Metrosideros

in mesic forest and more abundant in cloud

forest with Ile, v, Myrsine and Metrosideros up to

1200 m. On Tahiti, probably the most abundant

woody plant in undiscurbed forest from 400 m

to the summit of the island, Mt. Orohena, at

2241 m. Co-dominant wirh Metrosideros as the

main structural élément of slope and ridge forest

at medium and liigh élévations and practically

ubiquitous. Other associâtes in moss-covercd

shrubbery at high altitude in the typical

Weinmannia-Alstonta forest (Florence 1993)

include Myrsine , Coprosnui , Cyathea, Alstonia,

Vaccinium, Alyxiti, Heynoldsia , and //ex. At lower

élévations, ir occurs with Alphitonia , Dodonea

and Psidium and is also found in scrub, on

Dicranopteris- covered slopes and on steep wood-

ed escarpments.

SELECTED COLLECTIONS (from a total of 88 stu-

died).—S ociety ISLANDS: Bora Bora?-. Dumont

d'Urville 200, s.loc., fr. (P). Huahine: Florence &

Tahuaitu 11627, crête entre les Mrs. Mato Ereere et

Moua lu ri, l6°43VDrorW, 61(1 m, 11., 30 Oct.

1992 (P, PAP); Grant 5314, Distr. Maeva,

Matoereere. 1710 fr.. fl., 7 Peb. 1931 (BlSH); Grant

5316, seedling (BlSH); St. John 17159. Huahine Nui,

Mt. Matoereere. N ridge, 650 m- fr., 1 Ocr. 1934 (A,

BlSH, K- P). Moorea: Florence 7935, Haumi, crête

SE du Mt. Tohiea, P°33'S-149°49'W. 860 m. B., 22

Oct. 1986 (BlSH, K, P, PAP); Florence 8365, Mt.

Tohiea, sommet N, 17 Ù 33’S-149°49’W, 1200 m,

buds, 9 July 1987 (BlSH, P, PAP); Smith 174 , plat¬

eau above Afareairu, 11., 8 June 1967 (A, BlSH).

Tahiti: Adamson 40, Distr. of Hittaa, 400 ni, buds,

19 Nov. 1928 (BlSH); Carlquist 655, between Rocher

du Diable and Farc Mato on Aorai Mt., 4200 fî., fr.,

15 (une 1962 (GH); Florence 2291, sentier captage

d’eau du Belvédère, l7°34‘S-l49‘3rW, 620 m, fr.,

14 Jan. 1982 (P, PAP); Florence 2315, sentier du Mt.

Marau, au Pic Vert. 17°37'S-149”32'W, 1405 m, fl.,

fr., 28 |an. 1982 (BlSH, P, PAP); Florence 3277, Mt.

Marau, env, du relais TV, 1 7‘37’S-l49°32’W,

1430 m, yfr., 5 J une 1982 (BlSH, K, P, PAP);

Florence 3353 , route du Pîc Rouge, Propriété Lévy,

17“35’S-149“33 > W, 630 m, fr., 17 June 1982 (BlSH,

K. P, PA P); Florence 3645, créLe W de Ja Papenoo,

sentier de l’Orohena, 17 C 33'S-I49°26'W. 915 tn fr.,

3 Aug. 1982 (BlSH. K, P, PAP); Florence 5462,

Mahina. sommet du Mt. Pitohiti. sentier de

fOrohena, 17°37’S-149°28'\V, 2110 m, fr., 19 Oct.

1983 (P, PAP); Florence 7186, Mahina, sommer N de

1 Orohena, flanc S, I 7°37'S-149°29’W, 2215 m,

buds, 16 Nov. 1984 (P. PAP); Florence 9088, I aaa,

mute du Mt. Marau. km 6.5. 17°37'S-l49°34’W,

1080 m, fl., 11 Eeb. 1988 (BJSld, P. PAP); Florence &

Varney U003, Punaauia. vallée Maruapo, crête N,

17 a 39'S-l49"35'W, 880 m, fl., 13 Feb. 1992 (P,

PAP) ; Foshers 62927, N W slope up to Pic Vert, head-

waters of siae branches of R. Tipaerui, 640-650 m,

fr.. 17 June 1982 (BlSH. BM. K, P, PAP); Gagné &

Montgomery 2362, Mt. Orohena, 2240 m. 11., I 1 Sep.

1988 (BlSH); Gagné 1013 , Mt. Marau, 1493 m,

buds, fr., 30 June 1977 (BlSH, P); Grant 3601, Dist.

Punâauiâ, Diadem, 3275 Ft., fl., 14 May 1930

(BLSH); Hong!and & /-'lorener 12916. Faaâ, Mt.

Marau, ridge lovvards Pie Ven. belovv TV station,

1380 m. fl., 14 Mar. 1994 (MO, P, PAP); Hoogland

& Florence 12918, ridge between Upper laatautia

River and Paaraura valley, 600 m, fr., 16 Mar. 1994

(MO, P, PAP); Hoogland & Florence 12927 , Taiarapu

Peninsula, piateau de Taravao along ridge track in

upper Amoa catch ment, 680 m, fr., 17 Mar. 1994

(MO, P, PAP); Huugland & Florence 1293F between

Lac Vaihiria and Col Urufaau, 650 m, fl., 24 Mar.

1994 (MO, P. PAP); MacDaniels 1313. S side

Orohena, 1500 m, fl., 16 May 1927 (A. BlSH. K);

MacKee 3056. plateau below summit of Mt. Aorai,

1S00-2000 rn, fl., 27 Aug. 1955 (K); Moseley,

Challenger Expédition s.n., s.loc., 4000 ft.. Sep. 1875,

112

ADANSONIA, sêr. 3 - 1998 • 20(1)

Weinmannia in the central Pacific

(BM, K); Nadatud 413 , crêtes du Binai et du Tâfifï,

fl., Dec, 1856 (P); Quityle 39, Mt. Aurai Trail, !ea-

ward, 1694 m, fl., fr., 22 Sep. 1921 (B1SH, K);

Raynal & Taureau 16539, piste de T Aorai, entre

Belvédère et Pare Hamuta, 600 ni, buds, 6 Oct.

1971(P); Si. John O" Fqsberg 1/146, Orotena, S ridge,

1570 ni, fr., 25 Sep. 1934 (A, B1SH, P); Teraoka &

Kennedy 113, Papenoo, backofEric Garnier property,

fr., 6 Sep. 1979 (BTSH); van Balgooy 1790, NW ridge

ofAorai, 1700 tn, fl., fr., 22 Sep. 1971 (L).

Local NAM es.—A ito; Aito mou’a; Ouru. Local

uses: firewood (Quayle39).

TYPIFICATION. — Weinmannia parviflora was the

first species in the genus to be described front

this part of the Pacific and its taxonomie history

is disettssed by Bf.rnarD! (1964; 195). It was

described by G. FORSTFR, who with bis father,

was naturalist on the second of Captain Cook’s

circumnavigational voyages in the H.M.S.

Resolution frotn 1772 to 1775 (FOSBËRG 1993).

Accordingto MüRRIl.l. (1954: 206), the naine W.

parviflora was one of several that weee appropri-

ated by FORXTER frorn the unpublished manu-

script of SoLANLtER and for which the latter had

already prepared a detailed description based on

the BANKS & SoiANDER collections front the

first of Cook’s voyages. Llovvever, many of the

descriptions in ForsTLR’s Prodromus (1786)

were actually written during the second voyage

and D. NlCOLSON (pets, contm.), who is study-

ing the FORSI’ER types, considers it entirely

appropriate to choose one of the specimens in

the FORSTER collections as the lectotype.

The sheet selected at BA4 contains two élé¬

ments. The right Itand piece is designated as the

lectotype of Weinmannia parviflora and the left

hand piece belongs to W. racemosa L. f. front

New Zealànd. Both ol CoOKs expéditions went

on to collect in New Zealarid after visiting

Tahiti. Records of W. parviflora, mostly under

the name Leiospermum parvifforum, front New

Zealand [e.g. by D. Don (1830), WALPERS

(1846), G. Don (1834)1 are due to confusion

with W. race ni o sa. Both species Usually hâve

simple leaves of similar si/.e bur the inflorescence

structure is different (see HOPKINS 1998a),

Several early authors (e.g. GRAY (1854),

Mueller (1858), Seemann (1865), Drake

(1890)| also included material of Weinmannia

rapensis in their concept of W. parviflora. based

on Curning 1428. These records front Elizabeth

Island, (now Henderson Island in the Pitcairn

group) are discussed under W! rapensis.

VARIATION. — Weinmannia parviflora is especial-

ly variable in the shape of its leaves or leaflets,

but the young stents are usually pubescent and

the stipules strongly recurved and usually persis¬

tent. On Tahiti, W parviflora shows variation in

leaf shape with altitude, and our concept of this

species dilfers frorn BFRNARDl’s in including ail

the high altitude material with ovate or rounded

leaves here, rather than in W vescoi, which is

endemic to Raiatea, At middle élévations, the

internodes of W parviflora are relatively long and

rhe leaves are usually narrowly elliprical with the

margins markedly crenate or sinuate (Fig. IA),

and sortie triloliolate leaves occur (Fig. IL)). At

high élévation, in the central massif on Mt.

Aorai, Mt. Orohena and Pito Hiti, the leaves are

often shorter, usually ohlong or ovate and the

margins less markedly crenate (Fig. IB) and the

internodes are short, so that the overall habit

re.sembles that of W. vescoi■ Bçrween 1500-

1600 m on the flanks of Mt. Aorai, adjacent

trees can hâve contrasting leal morphologies

(J.C. BraDPORL» pers. comm,), Most collections

are densely pubescent on the young stents but a

few are glabrous (e.g. Gagné & Montgomery

2362 , Florence 7186; Florence 5464 is ± gla¬

brous).

Material front Pluahine, designated here as

Weinmannia parviflora , has sonie similarities

with W. raiateensis since the stipules are caducous

and the pctioles are winged when the leaves are

triloliolate. Collections front Moorea al.so dilfer

somewhat frorn W. parviflora in Tahiti as the

stems are more or less glabrous, the stipules are

rarely persistent and the leaves, altltough uni-

foliolate, hâve a rather sinuate and/or undulate

margin (Fig. 1C).

2. Weinmannia raiateensis J.W. Moore

Bernicc P. Bishop. Mus. Bull. 102: 29 (1933);

Bernardi, Bot. Jahrb. Syst. 83: 207 (1964) under W.

vitiensis. —Type: J.W. Moore 396, Society Islands,

ADANSONIA. sér. 3 • 1998 • 20(1)

113

Hopkins H.C.F. & Florence J.

Raiatea, on high ridge between Vairahi and Avera

Rahi vallcys, 300 m, 3 Dec. 1926 (holo-, BISH; iso-,

BISH!, P! 2 sheets).

Shrub or small tree 0.4-5 m high. Branching

sometirnes dichotomous. Nodcs scârcely thicken-

ed and annnlar scar not prominent; internodes

0.5-3.8 cm. Young stems sparsely strigose-

pubescent or glabrous. Stipules caducous or not,

elliptical, shortly spatludate to ligulate, some-

times recurved, 0.7-1-1 x 0.4-0.6 cm, apex

rounded to obtuse, ± glabrous witli short,

adpressed hairs towards base on abaxial surface.

Leaves mostly trifoliolate, sometirnes some uni-

foliolate, total leugth up to 9.7 cm (for a tri¬

foliolate leaf) irtcluding a pétiole of 0.4-2.4 cm;

pétiole semiterete and narrowly winged espectally

towards point of insertion of Ieaflets, glabrous or

rarely pubescent above or below; Ieaflets narrow¬

ly elliptical, elliptical or narrowly obovate, the

latéral oncs 2.3-6 X 0.7-1.7 cm, base cuneate and

unequal, apex acute, the terminal ones 3.8-8.3

(including a petiolule of 0.4-1.5) x 1.2-1.9 cm,

base long attenuate, apex usually acute, often

broken; the bladc glabrous on both sides, chartac-

eous to subcoriaceous, not punctate; margin

sometirnes rhick-ened, crenate, 9-14 notches on

each side ot a leaflet, midrib liât or slighdy raised

above, prominent below, secondary and tertiary

venation flat or slightly raised on both sides.

Inflorescence rather variable and nodes often

asymmetrically branched; eithet a central triad or

pentad, or two latéral triads, the apical bud abort-

ed; offen arising at nodes other rhan the most dis¬

tal on a shoot; peduncte and rachis segments

shortly and sometirnes densely pubescent,

peduncle 0.6-1.1 cm long, rachis segments up to

5 cm long. Floral buds inserted singly; floral

braers 0.6-0.7 mm long, scarcely longer than the

buds, ligulate, ± glabrous, caducous. Flowers uni-

sexual (male. Gagné 1457; female, Florence 8945);

pedicel 1-1.7 mm long; calyx lobes 0.6-0.7 mm

long, glabrous; petals 1.1-1.3 X 0.8 mm, oblong;

in tnale flower; filaments câ. 2.5 mm long, dise

lobes 0.4 mm long, ovary 0.7 mm long, pubesc¬

ent, styles 0.1 mm long, incurled; in female

Fig. 2.—Distribution of Weinmannia in the Society Islands.

114

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F. & Florence J.

flower: filaments 0.8-1 mm long, dise lobes

0.7 mm long, ovary 1-1.2 mm long, ± glabrous,

styles 0.9-1 mm long, the stigmas capitate.

Capsules 1.5-3.2 X 0.9-1.8 mm at dehiscence,

the styles ca. 0.5 mm long, exocarp glabrous;

calyx lobes caducous, central column présent.

Seeds 0.6 mm long, comose at both ends, the

hairs short, 0.4 mm, and not very abondant.—

Fig. 2, 3A-D.

JUVENILE LOUAGE, — Florence 3746b consists of

regrowth shoots Irom the base of the trunk. The

leaves are 3-5-foliolate, chartaceous, and the api¬

cal leaflet is almost rhombic and broader than in

the adult foliage.

BREEL1ING SYSTEM- —Limited herbarium mater-

ial has only unîsexual flowers, but this species is

in fact polygamodioecious (J.C. BkadfoRD pers.

comm.).

FlELD CHARACT1 US.— Leaves pale green.

Inflorescence rachis purplish; buds whîte or

cream; flowers pale yellow to white. Fruits red or

pale purple.

Distribution anp ecolocy. —Endémie to

Raiatea where it is relatively widespread but

rather scarce, from 300-965 m. A species typical

of hygrophilous végétation at mid altitude with

Metrosicleros, Glochidion and Myrsine , or in cloud

forest with l/ex, Macardngd and Astronidium. It

does not occur in the same habitats as

Weinmannia vescoi (sce below).

MaTEIUAL EXAMINEE).— SoCtETV JSLANDS: Raiatea.

Florence 3554, Plateau de Temehani Rahi, 16°47'S-

151”27’W, 690 m, fr„ 3 July 1982 (PAP); Florence

3746, ibid., 760 m, fr., 25 Àug. 1982 (BISFI, K, P,

PAP); Florence 3746b, ibid., regrowth, 25 Aug. 1982

(P, PAP); Florence 3754, ibid , flanc N\V. 740 m, fl ,

25 Aug. 1982 (P, PAP); Florence 5143, côte £, Avera,

Crête S de la Paipai, 16*48’S-! 51“26'W, 395 m, 13

Sep. 1983 (P, PAP); Florence 8945, Ievaitoa, crête

sommitale du Mc Toomaru, 16 o 50'S-151 "27 W,

965 m, fl., 27 Nov. 1987 (P. PAP); Gagné 1457b, Mt.

Temehani, NEsidc, 740 m, 3 Sep. 1977 (BISH, P).

Bernar i si (1964) placed Weinmannia raiateen-

sis in synonomy under W. vitiensis Seem. on the

basis of the type description. Both hâve trifoliol-

ate leaves but in VF raiateensis they are generally

smaller and more délicate, although there is some

overlap. Several other species in sect.

Leiüspernium in the Pacific hâve trifoliolate or

imparipinnatc leaves with a somewhat winged

petiole, including IL' crofiii , VF sp. (Solomons A)

and VF denhümii (Hopkins 1998c). Although

distinctions between taxa are slight, there are dif¬

férences in mean leaflet shape and dimensions.

This particular leaf morphology has thus cither

evolved several times or else thèse species origin¬

ale from a single trifoliolatc/impai ipinnate

ancestor that has been dispersed amongst a nam-

ber of volcanic islands and differentiated only

slightly on each.

3. Weinmannia vescoi Drake

III. Fl. 1ns. Pacif.: 35, t. 13 (1886); Fl. Polynésie

Franç.: 61 (1893); Bernardi, Bot. Jahrb. Syst. 83; 206

(1964);—Type; Vescoi s.n ., Tahiti (sic, but probably

Iront Raiatea) (Itolo-, P!).

Weinmannia ovalifolia J.W. Moore, Bish. Mus. Bull.

102: 28 (1933).—Type: Moore 95, Society Islands,

Raiatea, S facing slope along path to Mt. Temehani,

18 Sep. 1926 (holo-, BISH; iso-, BISH!, Ll).

Low shrub to small, multi-stemmed tree 0.15-

4 m high x 5+ cm diameter. Stems often sparsely

branchcd or unbranchcd lot long sections;

branching rarely dichotomous. Nodes thickencd

and “kneed", the internodes Iaterally contracted,

0.5-2.5(-7.5) longx ca. 1 mm thick. Stems glab¬

rous, smooth with fine longitudinal striations,

lenncels few, on lower parts of stems, Whole

plant largcJy glabrous except for sparse puberul-

ence sometintes présent on stipules, inflorescence

axes, pedicels; leaves, calyx and exocarp often

warty. Stipules usually caducous, elliptical, ligul-

ate or rarely orbicular, 0,3-0, 7 X 0.2-Ü.4 cm, apex

acute or rarely rounded, ± fiat or rarely recurved,

± glabrous to puberulent. Leaves unifoliolate,

(rarely trifoliolate and then the petiole winged),

the petiole 0.2-0.6 cm long, the blade elliptical,

broadly elliptical or ovate, 1,8-4.7(-6.4) x 1-

2.5(-5 5) cm, base trnncate to acute and shortly

decurrent, apex acute, obtuse or rounded, subcor-

iaceous to coriaceous, rarely punctate below,

margin sometimes thickened, crenate, crenatrons

more pronounced in smaller ovate or elliptical

leaves, 9-12 notches on each side ol each leaf,

midrib raised and ridged above, prominent

116

ADANSONIA, sér, 3 • 1998 • 20(1)

Weinmannia in the central Pacific

below, secondary venarion obscure, flar or slight-

ly indentcd above, slïghdy raised below.

Inflorescence usually a pentad sometimes with

rwo latéral triads, rarely a heptad, reduced leaves

sometimes présent at nodes witltin inflorescence;

peduncles and rachis segments glabrous or sparsc-

ly puberulenr, 0.5-1,2(-2.5) cm long, racemes

2.5- 5 cm long. Floral buds inserred singly; floral

bracts 0.7-1.4 mm long, ± eqttal in length to

buds or longer, ligulate to subeymbiform, minute-

ly ciliate on margtn. caducous. Flowers rnostly

unisexual, rarely bisexual; pedicel 1.4-2 1 mm,

calyx lobes 0.6-1 mm long; corolla 1-1.7 x 0.6-

1.2 mm, oblong; dise lobes 0.3-0.6 mm; in male

flowers: filaments 2.7-3.4 mm long, anthers 0.3-

0.4 mm diameter, ovary 0.6-1.1 mm, styles 0.1-

0.3 mm; in female flowers: filaments 0.7-1.2 mm

long, ovary 1-1.5 mm, styles 0.7-1 mm, stigmas

capitale, papillose; in bisexual flowers: filaments

1.5- 3.3 mm, ovary 1 2 mm, styles 0.7 mm.

Capsules 2.1 -4,0 X 1,5-1.8 at dehiscence, styles

0.5-1 mm long, stigmas capitate, exocarp glab¬

rous with longitudinal striations, endocarp some¬

times separating front exocarp; calyx lobes

caducous, central coluntn présent. Seeds ca.

0.8 mm long, comose at both ends, the hait s

0.3 mm long.-—Fig. 2, 3E-L.

BreedING SYSTEM. —Polygamodioecious. Most

specimens hâve male or female flowers (e.g.

Moore 793, St. John 172555, Florence 5197,

male; Florence 5203, 8991 and Hongland 12929,

female). Gagné 1454 bas some apparently

bisexual flowers, and Grant 5197 has mature

capsules and male flowers on the same shoot.

Field CH Alt ACIERS. —Growth form rarely with

a central trunk, but usually with numerous rail

stems arising from the base; often distinctive

with long, sparsely branched, srifl twigs with

short internodes and thickened nodes, the twigs

often well covered with unifoliolate, opposite

and decussate, elliptical to rounded leaves. Morat

6990 shows an extreme growdt form from dense

scrub in an exposed, wind swept région of the

plateau; ir is a much branched chamaepbyte, 15-

20 cm high with small, rounded leaves. In such

individuals the slender stems arise from a large

woody root-mass, often visible at ground level

(J.C. Bradford pers. comm.). Inflorescences

often short, congested, not projecting beyond

the leaves. Rachis of inflorescence and calyx

either green or purple-red; flowers similar to

those of W. parviflora in colour and scenr (J.C.

BRADFORD pers, comm.).

Distribution and ecology, —Endémie to

Raiatea where it is locally abundant from 355-

740 m. One of the species conftned to the plat¬

eaux ofTemehani Rahi and Temehani Ute Ute

where another endemic and symbol of rhe island,

Apetahia raiateensis (Campanulaceae), also

occurs. Locally co-dominant with Metrosideros,

Astronidium and Alstonia , the herb layer domin-

ated by Cyperaceae such as Gahnia seboenoides or

Machaerina bidwillii.

SriKCTED COLLECTIONS (from a total of 26 srud-

ied).— Society ISLANUS: Raiatea-. Florence 3586,

Plateau de Temehani Rahi, fin de la route, lôMô’S-

151°27’W, 490 m, fr., 5 July 1982 (BISH, P, PAP);

Florence 5197 , Plateau de Temehani Rahi, secteur NE,

16 u 46’S-151 ü 2è’W, 510 m, fl.. 16 Sep. 1983 (BISH,

P, PAP); Florence 8991, fevaitoa, sentier du Plateau

Temehani Ute Ute, IGM 7 T I51 1 '28\X/, 435 m, fl., fr.,

29 Nov. 198? (BISH, P, PAP); Florence 10)54,

fevaitoa, Plateau de Temehani Rahi. 16°46‘S-

I5D27'W, 530 m. fr., 10 June 1990 (BISH, PAP);

Garni 1457 (P). 1457a (BISH). Mt. Temehani, NE

siçfe, 740 m, buds. 3 Sep. 1977 (P, BISH); Grant

5234, Distr. Avcra. Temehani, 1690 ft., fl., 29 Jan.

1931 (BISH); idoogltnd & Florence 12929 , N slopes of

Plateau de Temehani rahi, 410 m. fr., 21 Max. 1994

(MO, P, PAP); Moore 793. path to Mt, Temehani, S

fa ci n g si ope, IL, 1 fan. 1927 (BISH); Morat 6993,

montée au Temehani uce, 550 m, buds, 26 Aug. 1982

(BISH, K, P, PAP); St. John 17255 , Temihani Plateau,

500 m, fl., fr., 5 Oct. 1934 (BISH, P); l Vbistler 4906,

ibid., 650 m. (1., 3 Sep. 1981 (BISH).

The type, Vescoi s.n., is labelled “Tahiti", The

lower parts of the stems are glabrous althottgh

the most distal inrernodes are moderarely hirsute

and the stipules are ligulate. The inflorescence is

not congested. The majority of collections from

high altitude on Tahiti that hâve similar broadly

ovatç leaves (Weinmannia parviflora) hâve thick-

er, more densely pubescent stems, strongly

recLtrvcd stipules and congested inflorescences.

However, a few have glabrous stems and ligulate

stipules and/or inflorescences which project

beyond rhe toliage. The idenrity and provenance

of Vescoi s.n. is thus problematical. it could either

be regarded as an aberrant form of W parviflora

ADANSONIA, sér. 3 • 1998 • 20(1)

117

Hopkins H.C.F. & Florence J.

from high altitude on Tahiti or treated, as here,

as a mislabelled collection from Raiatea.

Weinmannia i>eseoi resembles W. parviflora from

high altitude on Tahiti in sevcral respects, espe-

cially in the broadly ovate, oltcn coriaceous

leaves, the short internodes and thickened nodes.

However, it differs in having glabrous stems and

stipules that are usually caducous, ± fiat and

ligulate.

Although following the same synonomy as

BERNARD!, our cïrcumscription of this species

differs considerahly from bis, and St John 17255

from Raiatea is the only one of the specitnens

cited by BERNARD! that is referred here to

Weinmannia vescoi\ ail the others cited by him

are from Tahiti and placed by us in W. parviflora.

4. Weinmannia tnarquesana F. Br.

Bernice P, Bishop. Mus. Bull. 130: 99 (1935);

Bernardi, Bot. Jahrb. Syst. 83: 19.3 (1964).

a) Weinmannia marquesana var. marquesana

Weinmannia parviflora G, Forst. var. marquesana (F.

Br.) Fosberg in Fosberg & Sachet, Micronesica 8: 44

(1972).— weinmannia marquesana var. typica F. Br..

Bernicc P. Bishop. Mus. Bull. 130: 99 (1935), nom.

inval.—Type: Brown 497 , Marquesas Islands, Nuku

Hiva, To(o)vii. 1000 m, 15 )ulv 1921 (holo-, BISH;

iso-, B1SIP).

Weinmannia marques,ma F. Br. var. flabrata F. Br,,

Bernice P. Bishop. Mus. Bull. 1.30: 99 (1935).—

Weinmannia parviflora G. Forst. var. glabrata (F.

Br.) Fosberg in Fosberg & Sachet, Micronesica 8:

45 (1972).—Type: Brown 1084B , Marquesas

Islands, Hiva Oa, Mount Ootua, 800 m, 15 Dec.

1921 (holo-, BISH; iso-, BISFII).

Shrub to small tree, rarely a subshrub, (0.7-)

1.5-10 m tall, up to 35 cm dbh. Young stems

terete and narrow, 1-2 mm diameter, the nodes

slightly thickened, ± glabrous to densely coment-

ose (the hairs up to 0.5 mm long) or pilosc (hairs

up to 0.9 mm long), older stems glabrescent,

finely longitudinally fissured sometimes with

pale lenticels. Branching often dichotomous.

Stipules ustially early caducous, either elliptical-

ligulate, 0.6 X 0.2 mm the apex rounded and

recurved, or diamond-shaped to broadly ovate

and conduplicate 1.3-1.7 X 0.8-1.2 mm, the

apex acute and the margin entire to serrulate,

glabrous excepr for short strigose hairs on abaxial

surface towards the base. Leaves simple (rarely

trifoliolate), petiole semiterete, 0.3-1.8 cm long,

glabrous to tomentose along mid line on adaxial

surface and glabrous or more often pubescent on

abaxial surface, blade elliptical or sometimes

ovate, (2.5-)3.3-7.5(-9.6) x (l-)l,3-3,6(-4.4) cm,

base cuneate or acutely constricted into the

petiole, apex acute or sometimes acuminate,

blade subcoriaceous, glabrous, sometimes punct-

ate below, margin usually minurely thickened

and revolute, minutely crenulate, crenate, serrate

or rarely r ± sinuate, I 1-22 notches down each

side; main veiri fiat or raised ahove, and scime-

times minutely hairy towards the base, pro¬

minent and olten shortly hairy below, secondary

and tertiary venation fiat on both sides, the

secondary veins équidistant from one another

and ail arcuate towards the margin and apex at

the saine angle, réticulum of tertiary veins open

to dense.

Inflorescence a triad or pentad (variants include

3 triads or a central triad and 2 pairs of racemes

(each developing from a latéral auxiliary bud)

arising from the same node as the subtending

leaves), reduced leaves at nodes with in infloresc¬

ence not seen, pedunclex and rachis segments

0.8-2.5(-6.5) un long, almost glabruus or sparse-

ly to densely hairy, racemes up to 7-8(-12) cm

long. Floral buds inserted singly; floral bracts to

1.6 mm long, narrowly ligulate to subcarinate,

strigose-puberulent, caducous. Flowers unisexual

ot bisexual; pedicel 1-3.0 mm long, puberulent

or not; calyx lobes 0.7-1.2 x 0.6-1 mnt long,

shortly strigose or glabrous on outer surface;

petals oblong, 1.5-1.9 X 0.8-1.2 mm, apex round¬

ed; dise lobes 0.4-0.8 mm long; in male flowers:

filaments 2.7-3 mm long, the ovary 0.8-0.9 mm

long, usually pubescent, the styles ca. 0.2 mm

long, curvcd inwards; in female flowers: fila¬

ments 0.4-0.5 mm long, the ovary 1,1-1.5 mm

long at anthests, glabrous, and the styles 1 mm

long, ± erecr, tbe stigmas capitate and papillose,

ovules up to 4 x 7 per capsule; in bisexual flow¬

ers: filaments 2.8-3.5 mm long, ovary 1.5-

2.2 mm long, glabrous or with sparse short

strigose hairs, styles 1.2-1.4 mm long, stigmas

capitate, papillose. Infructescence often dense.

118

ADANSONIA, sér. 3 • 1998 • 20(1)

Hopkins H.C.F. & Florence J.

Capsules 3.3-5 x 1.5-2 mm at clehiscence,

styles 0.5-1 mm long, exocarp glabrous or almost

so, endocarp somedmes separaring from exocarp

in old fruits; calyx lobes caducous, central

column présent, Seeds 0.6-0.>3 mm long, comose

at each end, the hairs few and wealdy developed,

< 0.5 mm long.—Fig. 4A.C-G, 5.

JUVENILE LOUAGE. —Foliage from young plants

and coppice shoots is eithcr simple (Mumford &

Adamson 139 and Brown 1078) or triloliolate

(Mumford & Adamson 494) witli the stipules

often persistent. As in other species, the leaflets

are chartaceous or subcoriaccous. Perlman 10204

from low (otest on Hiva Oa i.s exoeptional as ir is

almost glabrous, has trifnliolate leaves up to

23 cm long (terminal leaflets up to 17 X 8.5 cm)

and the stipules are orbicular-reniform, 3.5 cm

diameter, with the apex hroadly rounded and the

margin entire or dentatc, This specimett is prob-

ably from a coppice shoot in shadc.

BREEDING SYS7 LM. — Polygamodioecious.

Several sheets hâve male flowers and mature

fruits and only 3 ouf of about 70 hâve female

flowers (Mumford & Adamson 497. Qudyle 1253

and Florence 7262). There is no évidence of pro-

tandry. Florence 7447 (P) has bisexual flowers

and dehisced fruits and another sheet (BISH) has

both male and bisexual flowers. This species

appears to hâve cycles of male and bisexual flow¬

ers, with occasional plants producing female flo¬

wers.

FlELD CHARACTHRS. —Variable in growth habit;

when a small tree, the crown sometimes umbrella-

shaped. Bark grey, brown or reddish, smooth;

sapwood white or pinkish; heartwood red, fla¬

grant. Flowers greenish-white to créant, calyx

greenish-red, corolla and stamens white. Young

fruits green to brigln carminé red.

Distribution and f.cology. —Marquesas

Islands; Paru Hiva, Hiva Oa, Nuku Hiva,

Tahuata, Ua Huka, and Ua Pou, from 500-

600 m upwards, except on Nuku Hiva where

found flom 790-1 180 m.

This appears to be a sister species of

Weinmannin punnjlora from the Society Islands

occupying the samc range of habitats, from dense

primary tain forest and disturbed forest at mid-

elevations to cloud forest and scrub on ridges

and slopes, often in exposcd localitiex, at high

altitude. It also occurs in xerophilous scrub or

open areas on hilltops with Dicranoptens, where

ir resprours after burning. lt is one of dte major

woody plants flom 790 m upwards on Nuku

Hiva, localfy co-dominant and tncreasingly com-

mon with altitude. In dry forest, scrub and

savanna its main associâtes are Metrosidrros and

Diannopteris , and also Paesia. In mesic forest it

occurs with Cyathea , Freycinetia , Cheirodendron ,

Santalum, Scaevola, Myrsine, llex, Glochidion,

Cyrtandra. Trimenia , Hernandia , Metrosideros ,

Mangifem, Pan dan us. Hibiscus , Wikstroemia , with

numerous ferns and epiphytes

SlJ.I.CTU) COI 1 l'.CnON.', (flom a rotai of 67 stud-

ied).— Marquesas: Fatu Hiva: Brown 1078, Omoa,

800 m, st., 4 Apr. 19l)5 (BISH); Decker 2388, along

trail between Omoa and Ui’a, withïn 1 km of pass

river central ridge, 750 m, fl., 12 Oct. 1974 (BISH,

P); Florence et al 9490 , haut bassin de l Uiha, E du

Mr. Teamotüa, KP2HS-1 3S*3S”W, 660 m, fl., 21

July 1988 (BISH, P, PAT); Gagne 1220. Teauapuhiau

Pass, above Ouia valley, 720 m, buds. 1-3 Aug. 1977

(BISH): Wagner et al. 6152. Ridge E of Mt.

7 earaorua flom ridge below road along ridge to casca¬

de, 640 m, fl., 2Î july 1988 (BISH). Hiva Oa :

llmwn 1084A, Feani, 900 m, st., 5 Apr. 1905 (BISH);

Cherner ih MacKet- 44711 , Mokoau, 500 rn, Ir., 13

Nov. 1989 (P, PAT’); Decker 1189 , Puatnau, along

Puamau-Atuona trail, 500-650 m, fl., 14 Dec. 1963

(L, P, P AP); Kondo s.n., Mt. la peata, 2500 fl,, fr., 15

Qcv. 1969 (BISH}; Mumford & Adamson 414, Mt.

Ootua, W slopc, 700 m. buds, fr., 25 May 1929

(BISH); Atumford dr Adamson 497, NW summit of

Mt. Temeriu, 3400 fl., fl., fl„ 3 Aug. 1929 (BISH);

Oliver & Schéifer 321 6, Mt. Ootua, old trail E of sum¬

mit, 640 m, fr., 26 Feb, 1975 (BISH, PAP); Perlman

10204, trail towards Hanamentt, 3200 fl., juv., 3

Aug. 1988 (B15I1, PAP); Perlman 10239, Vaipahee

Palis rd., 1700 fl., fr., 10 Aug. 1988 (BISH, K, P);

Sachet & Decker 1903, road flom Atuona to Puamau,

just below Mt. Ootua, 625-700 ni. fr., 23 Nov. 1974

(P, PAP). Nuku Hiva: Florence 4303, Tooviî, cpaule-

ment au-dessus du réservoir, 8°52’S-14 0”09’W,

950 ni, fr., 4 Dec. 1982 (BISH, P. PAP); Florence

8421, route Taiohae-Tnovii, branche droite de la Ht.

Taipivai, 8°5JS-140 0 08'W. 760 m, fl., 29 July 1987

(BISH, K, P, PAP); Florence 4361 , route Toovii-Terre

Déserte, km 6,8 après le col, 8°52’S-1 40°10’W,

960 ni, A., 9 Dec. 1982 (BISH, K, P. PAP); Florence

8338, Terre Déserte, crête Ouesr du Mt. Akaupe,

8°52'S-140“10’W, 1180 rn. buds, 10 Aug. 1987

(BISH, P, PAP); Gagné 1033, spur of Mt. Ooumu,

Toovii plateau, 790 m, fr., 16 July 1977 (BISH, L, P);

Gillett 2215, Toovii-Taiohae trail, 2 km from

120

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in the central Pacific

Tapuaooa sheltcr, 800 m, fr., 4 Aug. 1970 (BISH, K,

L, P); Lorence et al. 6089, Toovii région, NW of

l’Economie Rurale complex along new rd. to airport,

1060 m, fr., 16 July 1988 (P, PAP). Ta/mata: Halle

2168, sommet, fl., fr.. 17 Mar. 1973 (P); Périmait et

ai. 14911, summit of vidge above Vaicahu nr,

Haaoiputeomo, nr. antenna, 2740 fr., fl., 1 Sep. 1995

(P); Schtijer 5499, Vaitahu, crête d Amatea, 620 m,

st., 10 Apr, 1975 (K). Ua Huka. Florence 7262,

Hane, Crête SW menant an Mr. Hirikau, 8"55’S'

1.39°32’W, 540 m, fl., fr., 20 feb. 1986 (P, PAP);

Quayle 1679, .s.loc., fr.. *1 Nov. 1922 (BISH); Quayle

1758 (BISH); Quayle 1829, fl. (BISH); Quayle 1830,

fr. (BISH). lia Pou ; Quayle 1188, fr., 12 Sep. 1922

(BISH).

Local NAMES.— Atakua, Ata.

Distinction erom Weinaunnja I'arvihofa .—

Weinmannia marquesana and W. parviflora are

clearly closely related. Although BERNARDI

(1964) regarded W. marquesana as distinct,

FOSBERG & Sachet (1972) reduced it: to a varie-

ty of W parviflora ("table 1). We are once more

giving ir spécifie status although there is some

overlap in characters ( fable 2).

Variation within Weinmannia marques-

ANA .— Thts is a polymorphie species, varying in

the size and shape of the leaves and the quantity

and length of the indumentum, and thus it

shows much the same range in variation as

Weinmannia parviflora. Because of the range in

amount and type of indumentum, we agréé with

BERNARD! (1964) that W. marquesana var. ghb-

rata can not be mainrained. However, one parti-

cularly distinctive variant originally named by

FOSBERG & SACHET (1972) is maintained here,

var. myrsinites. Recuise of the variability of this

species, it is not surprising that there are some

intermédiares berween the two named varieties

(see below).

Florence 7262 is a specimen of Weinmannia

marquesana var. marquesana from Ua Huka with

female flowers, dehisced capsules and mostly tri-

foliolate leaves In W. parviflora on Tahiti, occa-

sional plants also occur with some rrifoliolate

leaves, either on the lower, stérile branches (e.g.

Hoogland & Florence 12911) or on flowering

branches (e.g. Hoogland & Florence 12915)-

Florence 6882 from 1166 m on Nuku Hiva lias

a mixture of long and short, relatively thick and

densely hairy internodes, and broadly elliprical

ot ovate leaves. It is thus vegerarively similar to

collections of Weinmannia parviflora from high

altitude on Tahiti. Périmait et al. 14911 from

Tahuata has exceptionally narrow leaflets.

Table 2.—Comparison of the characters of Weinmannia parviflora and W. marquesana var. marquesana

Character

W. parviflora

(at low and mid élévation)

W. marquesana var. marquesana

Shape of leaf blade

length-breadth ratio

narrowly elliptical

elliptical to ovate

excluding petiole

1 : (0.27-)0.3-0.37(-0.47)

1 : (0.36-)0.39-0.5(-0.58)

Stipules

persistent, recurved

elliptical, spathulate or ± orbicular

apex rounded, margln entire

caducous, oflen conduplicate

broadly ovate

apex acute, margin often toothed

(sometimes narrowly ligulate with

rounded apex and entire margin)

Capsules at dehiscence

2.5-3.5 mm long

3 3-5 mm long

Inflorescence

complex, racemes usually arising

from 3 successive nodes,

up to 11 racemes per inflorescence

(pentad + 6)

simple, racemes arising usually

at 1 node or sometimes 2,

triad or pentad (rarely otherwise)

Indumentum on young stems

dense indumentum of erect hairs

to 0.5 mm long

variable, dense to sparse

ADANSONIA, sér. 3 • 1998 • 20(1)

121

Hopkins H.C.F. & Florence J.

b. Weinmannia marquesana var. myrsinites

(Fosberg & Sachet) H.C. Hopkins & J. Florence,

comb. nov.

Weinmannia parviflora G. Forsr. var. myrsinites

Fosberg & Sachet, Micronesica 8: 45 (1972).—

Type: Sachet & Decker 1155 , Marquesas Islands,

Hiva Oa, Atuona-Feani trail, 1200-1300 m, 24-26

Sep. 1963 (holo-, US!; iso-, BJSH!, Kl. L, NY, P!).

Shrub 0.6-1.5 m high, less commonly a small

tree 2 m. Internodes usually short, ca. 0.5 cm

long, nodes thickened, young stems strigose to

puberulous. Leaves ovate or elliptical, the blade

1.6-2.7(-3.3) X 0.7-1-7 cm, plus a petioJe 1-

6 mm long, base cuneate or truncate, apex acute,

blade coriaceous, the upper side drying dark pur-

plish-brown, lower side dark chestnut, margin

with 8-12 notches on eirher side, venation usual¬

ly obscure and fiat above, secondary and rertiary

venation liai below and drying conspicuously

darker rhan intervenium, bases of secondary

veins frequentiy doser rogether in proximal part

of blade, and basal vein.s more strongly arched

towards apex than are the more distal ones, vena¬

tion ± brochidodromus, réticulum of tertiary

veins dense.—Fig. 4B, 5.

Fig. 5.—Distribution of Weinmannia in the Marquesas Islands.

122

ADANSONIA, sér. 3 ■ 1998 • 20(1)

Weinmannia in the central Pacific

FlELD CHARACTERS. —The inflorescences are

much shorter than in W. marcjuesdna var. mar-

cjuesana. Flowers white; young fruits red.

Distribution and ecolocy, —Marquesas:

Hiva Oa, 800-1300 m. Common in scrub on

upper, exposed, windswept slope at crest of ridge

(maquis sommital) on dry side ofisland and also

on leeward side in cloud forest.

Matériau eXAMIMED,— Marquesas: Hiva Oa.

Brown 1098, Action*, 800 m, sr„ Dec. 1921 (BISH);

Chemin in MacKee 41689. Feani, 1100 m, fl., tr., ] 1

Nov. 1989 (P. PAP); Florence & Périma» 9667,

Atuona, piste de Fl a n am cru, NW du Mt. Temetiu,

9 D 48’S-139°5’W, 1130 m, buds, Fr., 30 July 1988

(BISH, K, P, PAP): Oliver & Schÿfer 3138, Feani

ridge, 1150 m, tr., 12 Feb. 1975 (BISH, PAP); Oliver

& Schàfer 3148. ibid., 1050 m, Fr., 12 Feb. 1975

(BISH, P); Oliver é Schàfer 3155, ibid., 1020 m, Fr.,

12 Feb. 1975 (BISH, P, PAP); Périman 10198, trail

towards Hanamenu, 3550 ft., buds. 3 Aug. 1988

(BISH, P, PAP).

TaXONOMY AND VARIATION. —This variant of

Weinmannia manjuesana has a very distinctive

morphology, and sincc names are often useful for

such entities. wc retain it as an infraspecific

taxon. However, since it occurs only in more

exposed situations, it may perhaps be no more

than an ecotype of Weinmannia manjuesana. A

few collections from 1 liva Oa hâve leaves that are

either intermediate between the two varieties or

hâve characteristics ol both on same rwig ( Oliver

& Schafer 3161, Périma n 10230, Sachet &

Decker 1903, Decker 1189). Perlman 10130

(BISH, PAP) from Nuku Hiva has small leaves

on stems with thickcned inrernodes and dense

erect hairs. Its ven.nion is more similar to that of

W. marquesana var. mdrquesana than var. myrsin-

ites but the blades hâve dried almost black above

and dark chestnut below. It is placed here with

doubt.

5. Weinmannia tremuloides H.C. Hopkins &C

J. Florence, sp. nov.

Insignis huer omîtes Gallicae Polynesiae species laterali-

ter complanato petiolo; a W. raiareensi J.W. Moore

longiore petiolo, angtistioribus stipula, nutjoribus Jhicti-

bus, majoribus seminibus munitis longioribus pilis, prae-

cipue dijfert.

TYPE.— Florence & Perlman 9581 , Marquesas

Islands, Fatu Hiva, W ridge of Mt. Mounanui,

10"28’S-138“37’W, 700 m, buds, fr„ 26 July 1988

(holo-, BISH!; iso-, CHR!, P!, PAP!, PTBG!, US!).

Shrub ca. 2 m high. Branching sometimes

dichotomous. Stems finely longicudinally fissur-

ed, internodes 0.8-3 cm long, nodes somewhat

thickened. Plant almost cntircly glabrous, except

for buds wbich hâve strigose hairs ca. 0.5 mm

long, and occasional strigose hairs on young

stems, stipules and leaves. Stipules oblong to

ligulate. 0.6-0.8 X 0.2 cm, caducous. Leaves tri-

foliolate, total length up to 9 cm including péti¬

ole 2.5-4.2 cm; pétiole laterally compressed and

IJ-shaped in cross-section; leaflets narrowly elltp-

rical, the latéral ones 3.6-4.4 X 0.7-1 cm, sessile

and unequal at hase, apex acute; terminal ones

4.8-6 X 0.8-1.1 cm, base attenuate, apex acute;

the blade subcoriaceous, not punctate below, the

margin somewhat thickened, crenatc, 13-16

notches on each side of a latéral Icaflet; midrib

taised above, prominent below, secondary and

tertiary venation somewhat indented above and

Fiat below.

Inflorescence a central triad; peduncle and

rachis segments glabrous; peduncle 1-1.7 cm

long, rachis segments up to 6 cm long. Floral

buds inserted singly; floral bracts ca. 1.1 mm

long, ligulate, glabrous. Flowers at anthesis not

seen. Calyx in bud glabrous.

Capsule 4-5 mm at dehiscence, the styles ca.

1 mm, exocarp sparsely hairy or ± glabrous; calyx

lobes caducous, central column présent. Seeds ca.

0.9 mm long, comose ai both ends, the hairs

0.7 mm long.—Fig. 5, 6A-L.

Fl EU) CHARACTERS.— The foliage has a tremb-

littg appcarance because ol the long, laterally

compressed pétioles. Flower buds white.

Dehisced capsules brown.

Ecolocy and distribution. —Known ouly

from Fatu Hiva where it was found in low végé¬

tation on ridge top and cliffs with Metrosideros ,

Dicranopteris and Lycopodium, and described as

abondant from 700-850 m.

Para i y pic— M arquesas. Fatu Hiva-, Perlman &

Florence 10175, slopes of Mounanui above Vaieenui

Fails, 2300 ft., buds, 26 July 1988 (BISH).

ADANSONIA. sêr. 3 • 1998 • 20(1)

123

Weinmannia in the central Pacific

A well defined species. The leaflet shape is

reminiscent of some spccimens of Weinmannia

raiateensis but the petiole is quite different, lt is

the most glabrous species in the région and the

capsules are comparatively large, as in W mar-

quesana.

6. Weinmannia rapensis F. Br.

Bernice P. Bishop Mas. Bull. 130: 100 (1935);

Bernard!, Bot. Jahrb. Svst. 83: 201, t. 35 (1964).—

Type: J. Stokes 305, R.ipj, Kulukulu [Kuluku in pro-

tologue] or Onape, 140 ft., 13 Oct. 1921 (holo-,

BISH; iso-, BISH!).

Shrubby rree 1.5-6 tn high. up to 20 cm dbh.

Branchirig usually dichoromous witli central bud

aborted or il developed, then less strongly so

than latéral shoots. Young stems densely pubesc-

ent, older ones minutcly longitudinally fissured,

sometimes lenticellate. Buds in leal axils often

prominent. S'cipules usually caducous, rarely

seen, ca. 0.9 X 0.6 cm ar nnde subtending inflor¬

escence, ± glabrous on adaxial surface, strigose

on abaxial one, apex obtuse, up to 1 1 X 0.9 cm

on végétative shoots, margin toothed. Leaves

compound, leaflets usually 3 or sometimes 5;

petiole semiterete, 1-2.5 cm long, rachis seg¬

ments in imparïpinnate leaves 1.5-3 cm, petiole

and rachis densely pubescent on tlattened adaxial

surface; latéral leaflets elliptical to narrowly

ovate, 2.8-6.9 X 1.1-2.8 cm, base unequal to

dimidiate, apex acute; apical leallets elliptical or

obovate, 4.2-10.3 (including periolule up to

2 cm) X 1.7-3.5 cm, somewhar larger than latéral

leaflets, base often strongly attenuace to fornt a

petiolule, apex acute to acuminare; leaflet blades

glabrescenr except for midrib which is densely

pubescent above towards base and strigose belovv,

coriaceous, rarely puncta te below; margin rhick-

ened, sometimes undulate, markedly crenate,

crenations 14-26 clown each side and strongly

acroscopic; midrib sligbtly raised above and pro¬

minent below, secondary veins fiat or slightly

raised above and below.

Inflorescence eirber a central pentad, the lower

racemes subtended by fully developed leaves,

reduced leaves or neirher, or a pair of pentads or

triads, each developed from an axillary bud and

the apical bud aborted. Inflorescence rachis seg¬

ments 1-4 cm long, racemes up to 14 cm long.

Auxîliary buds rarely seen, probably aborred.

Stipules at nodes within inflorescence ovate,

caducous, smaller than sripules ar végétative

nodes. Floral buds inserted singly; floral bracts 1-

2 mm long, caducous. Flowers male, feniale and

bisexual; pedicçl ( 1.3-) 1.8-2,5 mm long, minute-

ly hairy; calyx lobes shorrly strigose or glabrous,

sometimes warry, 0 9-1 mm long; petals oblong,

1.6-1.9 X 0.8-1 mm; filaments in male and

bisexual flowers up ro 2.5-3.5 mm long, in fema-

le llower.s ca. 1.3-1.4 mm loug, not always of

equal length in any flower; dise lobes 0.5-

0.7 mm long; ovary in male flowers 0.7-0.9 mm

long plus styles 0.1 mm long, tn female flow'ers

1.5-1.7 mm plus styles 1.2-1.5 mm long (at stage

vvlien anthers still présent), variable in bisexual

flowers according to âge.

Capsules 2-3.5 mm at dehiscence, the styles

0.5-1 mm, exocarp glabrous with weak longitud¬

inal ridges, sometimes warry, endocarp some¬

times separating from che exocarp towards the

apex; calyx lobes persistent or not, central

column présent. Seeds 0.7-0.8 mm long, comose

at both ends.—Fig. 7A-E,

BrEEDING SYSTEM. —Polygamodioecious.

Different sbeets of 57. John & Fosberg 15304 bave

male flowers and young fruits. Halle 7646 lias

morphologically male flowers in one inflorescen¬

ce while another has bisexual ones. A. Stokes 357

and St. John cr Maireau 15531 both hâve old

female flowers.

FlEl.D CHARV I I-HS. —Bark rough, dark brtnvn,

sapwood reddish brown {St. John 15305). Leaves

bright green, coriaceous. Inflorescence rachis red,

fltnvers white; petals almost white, anthers flesh-

coloured, calyx and young capsules red; both

male and female flowers reporred as having a

faint fragrance like Ccanothus (St. John).

DISTRIBUTION. —Frohably endemic ro Rapa

island.

POSSIBLE OCCURRENCE IN THE PITCAIRN

GROUP.—BERNARDI (1964: 202) records this

species from the Pitcairn Group citing Cuming

1428 (K) from Elizabeth Island, tiow Henderson

Island, and Cumitig s.n. (G) from “Pitcairn (?)”.

There are two sheets at Kew labelled Cuming

ADANSONIA, sér. 3 • 1998 - 20(1)

125

Hopkins H.C.F. &c Florence J.

Fig. 7.— A-E, Weinmannia rapensis A. shoot with trifoliolate leaves, the stipules oaducous and inflorescence (1 of a pair ol latéral

triads with a pair of roduceo leaves at node); note latéral auxillary buds at bases of peduncles; B, male flower; C. female llower; D,

capsule at dehtscence: E. seed. (A. Florence 6514, B. D E. St- John & Fosberg 15304 ; C, Stokes 357. —F-M, Weinmannia raro-

tongensis F, shoot wïth simple and trifoliolate leaves, persistent stipules with rovolute margin and immature inflorescence (pentad);

additionai Inflorescences also developing In the exils of more proximal leaves; G detail of node showîng opposite pétioles and the

peduncles ol 3 partial inflorescences : noie scars close to the base of the two latéral peduncles and row of colletai s along stipular

scar; H, male flower; J, female flower after anthesis, most of the stamens fallen; K, immature fruit; L, capsule at dehiscence; M,

seed. (F, Merlin 306; G, K, Parks 22522\ H, MacKee 44309 ; J, Gardner 2503\ L, M, Wilder 543). —Drawn by C. Monnié.

126

ADANSONIA. sér. 3 • 1998 • 20(1)

Weinmannia in the central Pacific

1428. One gives the locaüty “Elizrh Island’ and

the other “1ns. Toubouia”. The record From

Elizabeth Island is doubtful For several reasons.

Firstly, accotding to St. JOHN (1940) Cuming

1428 was collected on Toubouia, now Tu b uai

Island in the Austral Group. Secondly, recent

extensive botanical collections on Henderson

Island, including the area where CUMING collect¬

ed, lailed to lind rhis species (Fl.ORF.NCE et al.

1995; S. WA1.DRF.N pers. comm.). Thirdly it is

known that some oF CuMINCs labels became

mixed up resulting in erroneous records (S.

WÀLDRFN pers. comm.). IF Cuming 1428 is From

Ttibuai, it is the first record From an island other

that Rapa; CUMING visited both Tubuai and

Rapa (St. John 1940: 88). The date ot collect¬

ion on one sheet is given as 1831 but 1828 is

considercd correct (Si. John 1940).

ECOLOGY. —From 30-220 m. Reported From

Forest on steep slopes and crest oF ridge, where

Metrosideros is dominant and also From

Dicranopteris-Metrosideros scrub on exposed

slopes. Other associâtes include Myoporum ,

Corokia and Fitchia. Sometimes very abundant

and Forming thickets [ILdlé 7646) but generally

less commun than is Weinmannia parviflora on

Tahiti. Rapa island is colder and wetter than the

Society and Marquesas Islands, silice it lies Far-

ther south and its climate is influenced by cool,

damp air currents From Antarctica (see PaSTUREL

1993).

Material examined. — Austral Islands: Rapa.

Cuming 1428, Rapa or Tubuai (K); Florence 6395,

crête E vers Mt. Morongouta, 27“37’S-144“21’W,

210 m, fl., fr., 2 Feb. 1984 (BISH, K, P, PAP);

Florence 6514, collines du NW de la baie de Haurei,

27°36'S-l44"2rW, 50 m. fl., 6 Feb. 1984 (BISH, P,

PAP); Florence 65/7, S de Haurei, êpaulemem N du

Mt. Maugaoa, 27“37’S-144“20’W, 75 ni, fl„ 7 Feb.

1984 (BISH, P, PAP): Huilé 7501, N vallée l liri,

150 m, st. v 2 Feb, 1984 (P); Flatté 7517 , Morugo-Uta,

200-220 m, m., 2 Feb, 1984 (P); Hat!/ 7646 , F du

fond de la baie d’Almrei, 30 ni, fl, f> Feb. 1984 (P);

Huilé 7700, N de File Karapoo Rahi, fr., 9 Feb. 1984

(P); Longfield 772, s.loc.. Il Apr. 1925 (BM, K);

Paulav 29. Maungaaiai, 250-350 m,22Jan. 1980 (A);

Raouls.n.. s.loc., fl. (P): A. Stokes 2, s.loc., 100 m, fl.,

6 June 1921 (BISH); A. Stokes357, Mauiigaaeac, 900

ft., fl., 19 Oct. 1921 (BISH); St. fohn & Fosberg

15304, Area, 75 m, fl., 1 July 1934 (A, BISH, L); St.

John & Fosberg 15305, ibid., fl., fr., 1 July 1934 (A,

BISH, K, L); St. John & Maireau 15531, Kainiam, S

ridge ofMt. Perahu, 400 ni, fl., 13 July 1934 (BTSH);

Varney 24, piste de Hiri, 35 m, fl., 3 Jan. 1990 (P).

Local NAME.—-Aito. Local uses: fîre wood and

canoë parts [A. Stokes 357).

Rather uniform and distinctive. Most similar to

Weinmannia rarotongensis (Fig. 7F-M), endentic

to Rarotonga, the chïef island of the Cook

Islands, from which it differs by having uniform-

ly trifoliolate leaves and being more velutinous

on the stems and pétioles. The branching is more

often diebotomous and the cienation of the leaf-

let margins are more acroscopic. The stipules are

usually caducous and not markedly recurved as

in W. rarotongensis.

Acknowlegdements

The work by HCFH was financed by the Human

Capital and Mobility Programme u( the Eutopean

Union via the nelwork on Boianic.il Diversity ol the

Indo-Pacific. Wc thank the herbaria cited for access to

specimens, the late R.D. Hoocn.ANn for his generosi-

ty in giving HCFH access to bis unpublished notes,

D Nu OISON for advice on the PORSTEfi collections,

P. LoWRY for his comntents on the manuscript, E.

ArmËDE for belp with typing, and C. MONNIÉ, D.

STORE Z and A. DeTTLOFF for the illustrations.

REFERENCES

BERNARD] L. 1964.—Revisio generis Weinmanniae.

Pars III: Sectiones III - IV - V -VI. Bot. Jahrb. Syst.

83: 126-184; 185-221.

Brousse R. 1993.—La géologie des îles hautes: t. 28-

30, in Allas de la Polynésie française. ORSTOM,

Paris.

Don D. 1830.—A monograph of the fàmily of plants

callcd Cunonîaceac. F.dinburgh New Philos. J. 9: 84-

96.

Don G. 1834.— A general history of the dichlamydeous

plants. London.

DrakE del CaSHi i o E. 1886-1892. — Ulustraiiones

flmae imu la ru m maris pacifie. Paris.

FLORENCE J. 1982. — Recherches botaniques en

Polynésie française. ORSTOM, Tahiti.

Florence J. 1993.—Végétation de quelques îles de

Polynésie Française : t. 54-55, in Atlas de la Polynésie

française. ORSTOM, Paris.

Florence J., Waldren S. & Chepstow-Lusty A.J.

ADANSONIA, sér 3 • 1998 • 20(1)

127

Hopkins H.C.F. & Florence J.

1995.—The llora of the Pitcairn Islands: a review.

Biol. J. Linn. Soc. 56: 79-! 19.

ForsïTR G. 1786 .—Florulae insularum Australium

Prodromus. Gôttingen.

FOSBERG F.R. 1993.—The Forster Pacific Island

Collections Ironi Captain Cook’s Resolution

Voyage. Allertmia 7: 41-86.

Fosberg F.R. & Sachet M-H. 1972.— Plants of

Southeastern Polynesta. 2. Micrunesiai 8: 43-49.

Gray A. 1854 .—United States Exp/orhtg Expédition,

vol. XV. Botany, Phanerogamia. New York.

Hopkins Il.C.F, 1998a.—A révision of Weinmannia

(Cunoniaccac) in Malesia and lhe Pacific. 1.

Introduction and an acconnt of die species of

Western Malesia, the Fesser Sonda Islands and the

Moluccas. Adansoma . sér. 3. 20: 5-41.

Hopkins H.C.F. 1998b.—A révision of Weinmannia

(Cunoniaccac) in Malesia and the Pacific. 2.

Suiawesi and the Philippines. Adansoma , sér. 3, 20:

43-66.

Hopkins H.C.F. 1998c.— A révision aï Weinmannia

(Cunoniaccae) in Malesia and the Pacific. 3. New

Guinea, Solomons Islands, Vanuatu and Fiji, with

notes on the species of Samoa, Rarotonga, New

Caledonin and New Zealand. Adansonia , sér. 3, 20:

67-106.

Merriu F..D. 1954. —The botany of Cook’s

Voyages. Chron. Bot. 14: 161-384.

MuEU.tR C.[K.] 1858.— Walper's Annales botanices

systématisât , vol. 5. Leipzig.

PaSTUREF I 1993,—Fa climatologie: t, 42, 43, in

Ados de ta Polynésie française. ORSTOM, Paris.

Sl.P, MANN B. 1865-1873.— Flora vitiemis. London.

St. John H. 1940. —Itinerary of Hugh Cuming in

Polynesia. Occas. Pap. Betnice Pauahi Bishop Mus.

16: 81-90.

WALPERS W.G. 1846.— Répertoriant botanices sys-

tematicae, vol. 5. Leipzig.

INDEX TO TAXA AND SYNONYMS OF WEINMANNIA

This index includes ail the taxa treated in the current révision of the Malesian-Pacific species (this paper and

HOPKINS 1998a,b,c, ail publishcd in Adansonia, sér 3, 20), thus some page numbers refer to the previous

papers. Accepted liâmes are in ordinary type, synonyms in italics, new naines in bold, and invalid names in

brackers. F = sect. Faseiadata , L = sect. Leiospermum.

Arnoldia jraxinifolia Blume ‘ Weinmannia fraxinea (D. Don) Miq.

helerophylbt Blume = Weinmannia fraxinea (D. Don) Miq.

pinnata Blume -- Weinmannia fraxinea (D. Don) Miq.

| Cortexpapeta>nn Runiph.) = Weinmannia fraxinea (D, Don) Miq.

Leiospermumparviflorum (G, Fotçt.) D, Don = Weinmannia parviflora G. Forst.

Marattist termina lis Sol. - Weinmannia parviflora G, Forst.

[Merretia terminalis Banks &r Sol, mis] Weinmannia parviflora G. Forst.

PterophylUfraxinea D. Don = Weinmannia fraxinea (D. Don) Miq.

Spiraeapinnata Blume = Weinmannia fraxinea (D. Don) Miq.

Weinmannia. p. 20

sect. Fasciculata Bentardi ex Hoogland & H.C. Hopkins, p. 21

sect. Leiospermum (D. Don) Engl., p. 21

[sect. Racemosae BernardiJ = sect. Leiospermum

affinis A. Grav L, p. 96

alta Engl. = Weinmannia fraxinea (D. Don) Miq.

aphanoneura Airy Shaw F p, 35

arnoldia A. Gray = Weinmannia fraxinea (D. Don) Miq.

blumei Planch. = Weinmannia fraxinea (D. Don) Miq.

var. major Ridl. = Weinmannia fraxinea (D. Don) Miq.

borneensis Engl. - Weinmannia fraxinea (D. Don) Miq.

[bulttsanensis Elnier.l = Weinmannia hutchinsonii Merr.

camiguinensis Limer = Weinmannia hutchinsonii Merr.

celebica Koord. F, p. 54

clemensiae Steenis F, p. 32

128

ADANSONIA, sér. 3 • 1998 • 20(1)

Weinmannia in the central Pacific

coodei H.C. Hopkins F, p. 52

croftii H.C. Hopkins L, p. 76

cuneatifolia Engl. = Weinmannia negrosensis Elmer

denharnii Seem. L, p. 86

dichotoma Brongn. 8c Gris L, p. 102

[dictyoneura Schltr. 1 = Weinmannia fraxinea (D. Don) Miq.

descombesiana Bernardi F, p. 44

devogelii H.C. Hopkins F, p. 48

duliternis Airy Shaw = Weinmannia fraxinea (D. Don) Miq.

exigua A.C. Srn. F. p. 83, 94

eymaeana H C. Hopkins F, p. 50

fraxinea (D. Don) Miq. F, p. 23, 69, 80

fraxinijblia (Blume) Miq. = Weinmannia fraxinea (D. Don) Miq.

furfuracea H.C. Hopkins F, p. 49

hooglandii H.C. Hopkins & J.C. Bradford F, p. 37

borsjieldii Miq. = Weinmannia fraxinea (D. Don) Miq.

hutchinsonii Merr. F, p. 56

hypoglduca Kaneh. & Hatus. = Weinmannia fraxinea (D. Don) Miq.

kajewskii Guillaumin = Weinmannia denhamii Seem.

[var. ambrymemh Guillaumin]

hypoghtued Kaneh. & Hatus = Weinmannia fraxinea (D. Don) Miq.

\irosinensis Elmer] = Weinmannia hutchinsonii Merr.

lucida Merr. F, p. 64

luzoniensis S. Vidal F, p. 58

var. puberula Elmer = Weinmannia luzoniensis S. Vidal

ledermannii Schltr. = Weinmannia fraxinea (D. Don) Miq.

maegillivrayi Seem. F, p. 85

manuana Christoph. L, p. 100

marquesana F. Br. L, p. 118

va r. glabrdtu F. Br. = Weinmannia marquesana F. Br. var. marquesana

var. marquesana

var. myrsinites (Fosberg &C Sachet) H.C. Hopkins & J. Florence, p. 122

[var. typica F. Br.| = Weinmannia marquesana F. Br. var. marquesana

monticnla DSniker - Weinmannia dichotoma Brongn. & Gris

negrosensis Elmer F, p. 55, 62

var. simplicifolm (Merr.) Bernardi - Weinmannia negrosensis Elmer

novoguineemis L.M. ferry = Weinmannia urdanctcnsis Elmer

ouaiemensis (Guillaumin fié Virot) Hoogland L, p. 102

ovalifolia J.W. Moore = Weinmannia vescoi Drake

paitensis Schltr. L, p. 102

papuetna Schltr. = Weinmannia fraxinea (D. Don) Miq,

parviflora G. Forst. L, p. 109

var. glabrata (F. Br.) Fosberg in Fosberg & Sachet = Weinmannia marquesana F. Br. var. marquesana

var. marquesana (F. Br.) Fosberg in Fosberg &C Sachet = Weinmannia marquesana F. Br. var.

marquesana

var. myrsinites Fosberg & Sachet = Weinmannia marquesana F. Br. var. myrsinites (Fosberg & Sachet)

H.C. Hopkins & J. Florence

var. pamiflom - Weinmannia parviflora G. Forst.

pullei Schltr. F, p. 70

purpurea L.M. Perry L, p. 83

racemosa L. f. L, p. 105

raiateensis J.W. Moore L. p. 113

rapensis F. Br. L, p. 125

rarotongensis Hemsl L, p. 100

rhodogyne Gibbs = Weinmannia richii A. Gray

richii A. Gray F, p. 92

samoensis A. Gray L, p. 100

serrata Brongn. & Gris L, p. 102

simplicifolia Merr. = Weinmannia negrosensis Elmer

ADANSONIA. sér. 3 • 1998 • 20(1)

129

Hopkins H.C.F. & Florence J.

spiraeoides A. Gray (doubtful species), p. 100

sylvicola Sol. ex A. Cunn. L, p. 105

sundana Miq. = Weinmannia fraxinea (D. Don) Miq.

tannaensis Guillaumin = Weinmannia denhamii Seem.

thomei Guillaumin = Weinmannia paitensis Schltr.

tomentella Schltr. = Weinmannia fraxinea (D. Don) Miq.

tremuloides H.C. Hopkins & J. Florence L, p. 123

trichophora L.M. Perry = Weinmannia urdanetensis Elmer

urdanetensis Elmer F, p. 65, 74

versteeghii L.M. Perry = Weinmannia pullei Schltr.

vescoi Drake L, p. 116

virgulata Schltr. = Weinmannia pullei Schltr.

vitiensis Seem. L, p. 98

ysabelensis L.M. Perry F, p. 82

sp. (New Guinea A), p. 79

sp. (Solomons A) L, p. 84

sp. (Solomons B) L, p. 84

sp. (Sulawesi A) F, p. 55

Windmannia blumei (Planch.) Kuntze = Weinmannia fraxinea (D. Don) Miq.

fraxinea (D. Don) Kuntze = Weinmannia fraxinea (D. Don) Miq.

horsfieldii (Miq.) Kuntze = Weinmannia fraxinea (D. Don) Miq.

sundana (Miq.) Kuntze = Weinmannia fraxinea (D. Don) Miq.

Manuscript received 30 October 1997;

revised version accepted 27 February 1998.

130

ADANSONIA, sér. 3 • 1998 • 20(1)

Landiopsis Capuron ex Bosser, genre nouveau

de Rubiaceae de Madagascar

Jean BOSSER

ORSTOM, Laboratoire de Phanérogamie, Muséum national d'Histoire naturelle,

16 rue Buffon, 75005 Paris, France.

Danielle LOBREAU-CALLEN

CNRS (UMS 826) et EPHE, Laboratoire de Phanérogamie,

Muséum national d'Histoire naturelle,

16 rue Buffon, 75005 Paris, France.

RÉSUMÉ

Description de Landiopsis Capuron ex Bosser, genre nouveau de Rubiaceae-

Cinchonoideae-Isertieae de Madagascar et de l’unique espèce connue :

L. caparonii Bosser. L’étude palynologique confirme ses affinités avec le genre

Mnssaenda.

ABSTRACT

Description of Landiopsis Capuron ex Bosser, a new genus of Rubiaceae-

Cinchonoideae-lsertieae from Madagascar and of L. capuronii Bosser, the

only known species. Palynological study confirms its affinities with the genus

Mussaenda.

CAPURON, à son décès en 1971, avait légué

toutes ses notes et documents à Mme Françoise

Chauvet (Capuron 1973). En ce qui concerne

les Rubiaceae, les notes furent regroupées et dac¬

tylographiées en un seul document par Mme

CHAUVET qui demanda à l’un de nous (J.

BOSSER) de les revoir, CAPURON avait travaillé sur

cette famille de longue date et il était évident

qu’il n’aurait pas lui-même publié cette étude en

l’état. Il fut donc décidé, en accord avec Mme

CHAUVET, de tirer ce document en plusieurs

exemplaires, afin que les chercheurs travaillant

sur les Rubiaceae de Madagascar puissent y avoir

accès. Ils furent déposés au CTFT (Paris,

Tananarive) et un exemplaire fut confié au

Professeur J. -F. LEROY du Laboratoire de

Phanérogamie, Muséum de Paris. Ce dernier se

chargea de l’étude de certains genres

( Mantalania, PseudomantaLinia ) mais ne se pen¬

cha pas sur le cas du genre Landiopsis, proposé

aussi par CAPURON, dont il nous remit récem¬

ment le matériel.

MOTS CLÉS

Rubiaceae,

Landiopsis,

palynologie,

nouveau genre,

Madagascar.

KEYWORDS

Rubiaceae,

Landiopsis,

palynology,

new genus,

Madagascar.

ADANSONIA, sér, 3 • 1998 • 20(1)

131

Bosser J. & Lobreau-Callen D.

Il s’agit d’un arbuste du NW de Madagascar,

sans raphides dans les tissus, à stipules inierpécio-

laires bifides , inflorescences terminales

cymeuses ; fleurs grandes ; calice à 5 lobes subu-

lés ou, sur certaines fleurs, un des lobes dévelop¬

pé en lame foliacée ; corolle longuement

tubuleuse, à 5 lobes imbriqués ; étamines 5,

incluses, subsessiles ; ovaire biloculaire, à 2 pla¬

centas axiles, multj-ovulés ; style court, à 2

branches stigmatiques cylindriques (des fleurs

longistyles n’ont pas été observées) ; fruit capsu¬

laire, déhiscent ; graines nombreuses, irrégulières,

non ailées, finement ponctuées-réticulées.

LANDIOPS1S Capuron ex Bosser, gen. nov.

Mussaendae Burm. f. atque Pseudomussaendae

Werhn. afjinis ; a primo aestivatione imbricttta etfnictu

capsulari, a secundo aestivatione imbricata, praecipue

differt.

Type. — Landiopsïs capuronii Bosser.

Le nom Landiopsïs est basé sur Landia proposé

par CoMMERSON et repris par PoiRET pour un

Mussaenda des Mascareignes.

Landiopsïs capuronii Bosser, sp. nov.

Frutex sine raphidibus, 4-5 metralis. Inflorescentiae

terminales, cymosae. Flores 5-meri, magni. Calycis lobis

linearibus. uno aliquando faliaceo. Corolla lobispatenti-

bus, imbricaris ; tuba cylindrico, Stamitia 5, subsessilia ,

inclusa, infra corollae orem affixa. Ovarium biloculare ;

placentis tixialibus. midtiovuLitisque ; Stylus incluais,

brevisque, stamitia hatid attingens. Discus anmdaris.

Capsula septicidalis . Sembla multa min iliaque, plusmi-

nusve renifbrtnia. rxdlntn ; testa subtiiiter cetiouldta .

Type, — Service Forestier 24467-SF, Capuron,

Madagascar, 3 fév. 1966 (holo-, P ; iso-, B, BR, G, K,

MO, P, PRE, TEF, WAG).

Arbuste atteignant 4-5 m de hauteur, sans

raphides dans les tissus ; rameaux jeunes densé¬

ment pileux, poils hirsutes, atteignant 1 mm de

longueur ; rameaux âgés ± glabrescents, bru¬

nâtres, à ienticelles allongées plus claires. Feuilles

opposées, caduques, membraneuses, discolores, à

face inférieure plus pâle ; limbe plan, entier, ovale

ou elliptique à largement ovale ou largement

elliptique, cuspidé au sommet, cunéiforme à la

base, un peu dissymétrique et décurrent sur le

pétiole, 5-10 X 2,5-6 cm ; faces supérieure et

inférieure pubescentes, à poils situés sur les ner¬

vures et les nervilles ; réseau des nervilles noi¬

râtre, ressortant sur le fond plus clair ; pétioles

grêles, densément pubescents, longs de 6-

18 mm. Stipules interpétiolaires, longues de 6-

7 mm, bifides, chaque lobe prolongé en arête très

fragile, pubescentes sur les 2 faces, finalement

fendues jusqu’à la base, ayant à leur aisselle une

ligne très dense de poils blancs. Inflorescences

terminales, 3-15-flores, en cymes dichotomes

assez denses, sessiles, les 2 premières ramifica¬

tions partant de la base ; cymules pédonculées,

les pédoncules densément pubescents, longs de ±

5 mm ; bractéoles 2, longues de * 1 cm, linéaires,

parfois rri-aristées avec 2 arêtes latérales beau¬

coup plus courtes, ayant à leur aiselle un groupe

d'arêtes plus courtes représentant une fleur avor¬

tée. Fleurs grandes, blanches, 5-mères, sessiles ou

courtement pédicellées. Calice à 5 lobes subulés

pubescents-hirsutes, un peu soudés à la base,

longs de 0,7-1,8 cm ou, plus souvent, l des lobes

beaucoup plus grand et développé en lame folia¬

cée, pétiolée, blanche, pubescente puis ± glabres-

cente, lancéolée ou étroitement elliptique, aiguë,

2-3 x 0,8-1,2 cm, devenant scarieüse sur le fruit,

à pétiole long de ± 1,5 cm, pubescent. Corolle

tubuleuse, longue de 5-6,5 cm ; tube long de

3,5-5 cm, cylindrique sur les 2/3 inférieurs et de

2-2,5 mm de diamètre, dilaté au niveau des

anthères et de 4 mm de diamètre, extérieurement

à pubescence dense, antrorse, apprimée, blanche,

intérieurement à pubescence hérissée, clairsemée,

.sous la zone d’insertion des étamines ; lobes 5,

imbriqués, lancéolés ou étroitement elliptiques,

aigus, finalement étalés, 10-12 x 5-7 mm, pubes-

cents sur les 2 laces, les nervures médianes

saillantes à la base du cAté interne, se rejoignant

et fermant la gorge. Etamines 5, incluses, insérées

au 1/4 supérieur du tube de la corolle ; anthères

étroitement oblongues, apiculées au sommet,

bilobulées à la base, vert olive, à déhiscence

introrse, subbasifixes, subsessiles l’extrémité du

filet ± soudé au tube, longues de 4,5-5 mm.

Pollen en grains simples tricolporé. Ovaire obeo-

nique, environ 5x2 mm, pubescent-blanchâtre,

biloculaire, à placentation axile, chaque loge

132

ADANSONIA, sér 3 • 1998 • 20(1)

Bosser J. & Lobreau-Callen D.

contenant un placenta allongé fixé sur toute sa

longueur ; en coupe transversale, placentas en

forme de T, les ovules étant fixés aux extrémités

des branches du T ; ovules très nombreux, hori¬

zontaux ; style inclus, long de 2,5-3 cm, glabre,

divisé au sommet en 2 branches stigmatiques

longues de ± 2,5 mm, apiculées» pubescentes sur

le dos et papilleuses sur la face interne ; stigmate

situé sous la zone d’insertion des anthères ;

disque présent, haut d environ 0,5 mm, entou¬

rant la base du style. Fruit, capsule à déhiscence

septicide, ovoïde ou ellipsoïde, 8-10 X 5-6 mm,

pubescente, faiblement côtelée, portant quelques

lenticelles blanchâtres saillantes ; calice persistant

mais non accrescent. La capsule finalement se

déchirant en 4 valves, l’exocarpe se détachant de

l’endocarpe. Graines nombreuses, très petites,

d’environ 1,5-2 X 1 mm, non ailées, + réni-

formes, irrégulières, comprimées latéralement, à

tégument finement réticulé. — Fig. 1.

Espèce de la forêt décidue sèche du Nord-Ouest

de Madagascar, rare, connue seulement par 2

récoltes.

Matériel étudié. •— Madagascar : Service

Forestier 24467-SF, Capuron , NW, massif de

l’Ankotekona, au S de la baie d’Ambararata (Diego-

Suarez), ait, 150 m, 3 fev. 1966, fl., type (B, BR, G.

K, MCI, P, PRE. TEL, WAC) ; Service Forestier

24668-SF, Capuron, même localité, entre 25 et 265 ni

d’alt., 25 avr. 1966, fr. (BR, G, K, MO, P, TF.F,

WAG).

Par le port, les grandes fleurs, les caractères

séminaux, cette plante rappelle certains

Mussaenda (Isertieae) comme le M. landia Poir.

des îles Mascareignes. Cependant, l’association

des 2 caractères : corolle à préfloraison imbriquée

et fruit capsulaire déhiscent, semblait la rappro¬

cher de la tribu des Rondeletieae.

Pour tenter de préciser la position de ce genre,

l’un de nous (D. LOBREAU-CALLEN) en a entre¬

pris l'étude palynologique.

Morphologie du pollen

Le pollen des Rubiaceae a fait l’objet de nom¬

breuses publications (Thanikaimoni 1972-

1986 ; Tissol 1990 ; Tissot Sc Van df.r Ham

1994). Dans plusieurs espèces, il présente un

caractère apertural rare chez les Eudicotylédones

(LobreAU-CaI.LHN 1978) : l'aperture composée

des grains colporés ou pororés est constituée de 3

parties, une ectoaperture avec une membrane

souvent granuleuse, une mésoaperrure circulaire

entourée d’un annulus de foot-layer et une

endoaperture circulaire ou elliptique située dans

l'endexine (LOBREAU-C AMEN 1978 ; VASANTHY

1978 ; ANfiERSSON & PERSSON 1991 ; PERSSON

199.3 ; Pufe et al. 1993 ; Huysmans ét al. 1994).

Ce type pollinique est dérivé d'un pollen tricol-

poré formé de 2 pairies, une ecto- et une endo¬

aperture (LoBREAU-CaI I EN 1978). Le grand

développement de l’annulus et l’augmentation

du nombre des apertures sont aussi considérés

comme des caractères évolués (RoBBRFCHT &

PUFE 1986 ; ANDERSSON & PERSSON 1991).

l,e pollen de l.andinpsis a été comparé au pollen

d’espèces appartenant à la tribu des Isertieae :

Isertia cacchiea (Aubl.) GmeL [de Granville 64,

Arataye, Guyane (P)] ; Mussaenda frondosa L.

[ Balama 625 , Tonkin, Viêt-Nam (P)) ; M. lan¬

dia Poir. [ Friedmctnn 463. La Réunion (P)] ; et à

la tribu des Rondeletieae : Rondeletia bourgaei

Standl. [Botteri 1016 , Mexique (P)] ; R. budd-

leioides Benth. [Gitleotti 2682H, Yotao, Mexique

(P)l-

Les pollens ont été acétolysés et observés au

Mph sur un appareil Zeiss standard et au MeB

sur un microscope Jéol 840A après métallisation

à for-palladium.

Le pollen de Landiopsis (Fig. 2A-D) est bré-

viaxe, isopolaire et brévi-tricolporé (Fig. 2A).

L’aperture est complexe et constituée de 3

parties :

— une ectoaperture large, courte et arrondie aux

extrémités, plus développée vers l’un des

pôles et recouverte d'une membrane scabre

(Fig. 2B) ;

— une mésoaperture circulaire, entourée d’un

annulus (Fig. 2C) ;

— une endoaperture elliptique, allongée sur

l'équateur (Fig. 2C).

L’exine massive (Fig. 2D) est formée par un tec-

134

ADANSONIA, sér. 3 • 1998 • 20(1)

Landiopsis , nouveau genre de Rubiaceae de Madagascar

Fig. 2. — A D. Landiopsis capuronii , E-H, Mussaenda landia : I-K. M. frondosa : L N Isertia coccinea O-P, Rondeletia bour-

gaei ; Q-R, R. buddleioides . A, E, L. O. Q, pollen trîcolporé ; B, F, J, ectoaperture ; C, G. endoaperture et mésoapedure entourée

d'un annulus ; D, H, structure massive de i'exine ; I, pollen tétracolporé ; K, exme avec de très courtes columelles ; M, infratectum

columellaire et nexine endocraquelée ; N, P, endoaperture subrectangulaire et nexine endocraquelée ; R, infratectum columellaire et

nexine endocraquelée. (Échelle : A-D, F-K, M, O-R = 1 pm ; E, L, N = 10 pm).

ADANSONIA, sér. 3 ■ 199B • 20(1)

135

Bosser J. & Lobreau-Callen D.

Tableau 1 . — Comparaison des caractères polliniques de Landiopsis et des trois genres proches.

Caractères polliniques

Landiopsis

Mussaenda

Isertia

Rondeletia

Symétrie

± isopolaire

isopolaire

Apertures (nombre)

3-4

Ectoaperture

courte, large, parfois

dissymétrique

longue, symétrique

ou non, parfois

à peine distincte

courte et étroite,

parfois

dissymétrique

longue et

étroite

extrémités

arrondies

aiguës ou dichotomiques

aiguës

membrane

lisse ou scabre

granuleuse

Mésoaperture

circulaire

absente

annulus

net

absent

Endoaperture

elliptique

circulaire

subrectangulaire

Tectum

lisse

perforé ou fovéolé

perforé

Infratectum

indistinct

columellaire ou indistinct

columellaire

Nexine

micro-endosculptée

micro-endosculpfée

endocraquelée

tum lisse, un infratectum de structure indistincte

au Mph comme au MeB, et une nexine finement

endosculptée.

Par ses caractères macromorphologiques, le pol¬

len de Landiopsis a des affinités avec celui des

Isertieae et des Rondeletieae. Chez les Isertieae,

Puff et al. (1993), AnderSSON (1996) et

HuYSMANS et al. (coram. pers.) ont étudié le pol¬

len de l’ensemble des genres du groupe. Les

grains sont 3(-4) colporés et le tectum est perfo¬

ré, « fovéolé », ou réticulé ; l’infratectum est colu¬

mellaire, rarement de structure indistincte

(Isertia p.p.). Le pollen d ’ Isertia diffère de celui

des autres genres par son sillon court et relative¬

ment indistinct alors que celui de Mussaenda se

caractérise par un sillon généralement tronqué

aux extrémités et par un annulus autour de

Pendoaperture. Le pollen des Rondeletieae est

tricolporé et le tectum nettement perforé

(BoRHini 1982 ; Borhidi & Jarai-Komlodi

1983 ; IgeRSHëim 1993).

Dans l’étude que nous avons faite, seul le pollen

de Mussaenda tri- ou tétracolporé (Pig. 2E,I) pré¬

sente une aperture formée de 3 parties dont une

ectoaperture longue (Fig. 2F,J), une mésoaperru-

re circulaire entourée d'un annulus (Fig. 2G) et

une endoaperture concentrique. Le pollen de

Mussaenda rappelle donc celui de Landiopsis.

Cependant, chez Mussaenda, la membrane aper-

turale est granuleuse (Fig. 2F,J) alors qu elle est

scabre chez Landiopsis. En revanche, dans les

deux genres Isertia et Rondeletia , le pollen est tri¬

colporé (Fig. 2L,0,Q) et l’aperture ne comprend

que 2 parties : une ectoaperture allongée, courte

lorsque les grains sont bréviaxes ( Isertia. , Fig. 2L)

ou longue lorsqu’ils sont longiaxes ( Rondeletia )

et, dans tous les cas, une endoaperture subrectan-

gulairc (Fig. 2N,P).

Dans tous les genres, le pollen est tecté, réguliè¬

rement perforé, parfois finement ( Isertia,

Mussaenda) (Fig. 1Î,L,0,Q) et l’inftatectum est

distinctement columellaire (Fig. 1K,M,R). Plus

rarement, le tectum est lisse, à peine perforé et

fossulé (Fig. E) et la structure de finlratectum est

indistincte (Fig. 1H). Ainsi, par les caractères de

la sexine, le pollen de Landiopsis montre des affi¬

nités avec ceux de Mussaenda et d'Isertia.

Chez les Rubiaceae, la nexine peut être lisse,

endosculptée ou moins fréquemment cndocra-

quelée et alors formée de plaques dans sa partie

profonde (AhABIE & KLPDAM-MaI.FRANCHE

1975 ; HUYSMANS et al., comm. pers.). Ce carac¬

tère se retrouve chez Isertia et Rondeletia où il est

particulièrement net (Fig. 2M,N,P,R). En

revanche, cette même couche est micro-endos-

culprée chez Mussaenda (Fig. 2H,K) comme chez

Landiopsis.

En conclusion (Tableau 1), le pollen de

Landiopsis est original par sa membrane apertu-

rale lisse. Cependant, il présente des affinités évi¬

dentes avec celui de Mussaenda puisqu’ils ont en

commun le même système apcrtural et la nexine

micro-endosculptée.

136

ADANSONIA. sér. 3 • 1998 • 20(1)

Landiopsis , nouveau genre de Rubiaceae de Madagascar

Par les caractères principaux des stipules, de

l’inflorescence, de la fleur, du fruit et de la graine,

le genre Landiopsis fait partie de la tribu des

Isertieae telle qu elle a été circonscrite par

Andersson (1996),

Remerciements

Les auteurs remercient Monsieur le Professeur J. -F.

LEROY qui leur a confié cette étude ; Madame N.

D’Amico (EPH E) responsable des techniques de paly¬

nologie ; Madame J. LEMEUX, auteur de la planche ;

Monsieur J. FLORENCE (ORSTOM) qui a traduit les

diagnoses latines. Les observations au McB ont été

effectuées au Centre Inter-universitaire de

Microscopie électronique de l’Université Pierre el

Marie Curie de Paris avec le concours technique de

M. Grasse et de F. Devienne. Les tirages photos ont

été réalisés par P. LOBREAU, S, HUYSMANS & E.

ROBBRECHT ont bien voulu nous faire part de leurs

travaux les plus récents sur la tribu des Isertieae.

RÉFÉRENCES

Abadie M. & Kl ddam-Maiblanchi M. 1975. —

Étude au microscope électronique à transmission du

sporoderme de deux espèces de Rubiaceae. Bull. Soc.

Bot. France, Coll. Palynologie, î 22 89-91.

Andersson L. 1996. — Circuinscription of the rrihe

Isertieae (Rubiaceae). Opéra Bot. Be/g. 7 : 139-164.

Andersson L. & Persson C. 1991. —

Circuniscrlption of the tribe Cinchoneae

(Rubiaceae) — a cladlstic approach. PI. Syst. Evol.

178:65-94.

BORHIDI A. 1982. — Studies in Rondeletieae

(Rubiaceae) 111. -— The généra Rogiera and

Arachnothrix. Acta Bot. Acad. Sci. Hung. 28 : 65-72.

Borhidi A. & Jarai-Komlodi M, 1983. — Studies

in Rondeletieae (Rubiaceae) IV. A new genus

Javorkaea. Acta Bot. Acad. Sci. Hung. 29 : 13-27.

CapüRON R. 1973. — Révision des Rubiacées de

Madagascar et des Comores. Manuscrit déposé au

MNHN, Paris.

Huysmans S,, Robrrecht F„ & SMETS F,. 1994. —

Are the généra Hallea and Mitragyna (Rubiaceae-

Coptosapelteae) pollen morphologically distinct ?

Blurnea 39 : 321 -340.

klERSHEIM A. 1993. — The paiynology of the genus

Rondeletia 1 . (Rubiaccae-Cinchonoideae-

Rondeletieae). Gratta 32 : 321-326.

LoiSRfiAU-CALLEN D. 1 978. — L’aperture composée

des Rubiaceae. Ann. Mines Belg. 2 : 167-173.

Persson C. 1993. — Pollen morphology of the

Gardenieae-Gardeniinae (Rubiaceae). Nord. J. Bot.

13: 561-582

PUFF C., IGERSIO IM A, & ROHRHOFFR U. 1993. —

Pseudotnussuenda and Scbizonutssaenda (Rubiaceae) :

close allies of Mussaenda. Bull. Jard. Bot. Belg. 62 :

35-68.

Robrrecht E. & Puff C. 1986. — A survey of the

Gardaiieae and rclated tribes (Rubiaceae). Bot.

Jabrb. Syst. 108 : 63-137.

ThANIKAIMONI G. 1972-1986. — Index bibliogra¬

phique sur la morphologie des pollens d'Angiospermes.

lnstit. Français de Pondichéry, Trav. Scct Sci, Tech.

12 (1), 1972 ; 12 (2), 1973 , 13, 1976 17, 1980 ;

22(1986).

Tissot C. 1990. — Sixième index bibliographique sur

la morphologie des pollens d'Angiospermes. lnstit.

Français de Pondichéry, Trav. Secr.. Sci. l’ech. 27.

Tissot C. & Van der Ham R.W.J.M. 1994.—

Septième index bibliographique sur la morphologie des

pollens d'Angiopermes. Insut. Français de Pondichéry,

Publ. Dept. Ecologie 36.

Vasanthy G. 1978. - Complexifies of aperrures,

columellae and tectum. IV Intern. Pdlyrtol. Conf.

Lucknow (1976-1978) : 222-227.

Verdcourt B. 1958. — Remarks on the classifica¬

tion of the Rubiaceae. Bull. Jard. Bot. Etat 28 : 209-

281.

Manuscrit reçu le 12janvier 1998 ;

version révisée acceptée le 13 mars 1998.

ADANSONIA, sér. 3 • 1998 • 20(1)

137

A floristic inventory and preliminary végétation

classification of the mixed semi-evergreen rain

forest in the Minkébé région, North East Gabon

Johan L.C.H. van VALKENBURG

Department of Plant Taxonomy, Herbarium Vadense,

Wageningen Agricultural University, P.O Box 8010,

6700 ED Wageningen, the Netherlands.

johan.vanvalkenburg@prosea.pt.wau.nl

Pieter KETNER

Department of Terrestrial Ecology and Nature Conservation,

Wageningen Agricultural University, Bomsesteeg 69,

6708 PD Wageningen, the Netherlands.

pieter.ketner@staf.ton.wau.nl

Chris M. WILKS

Africa Forest, B.P. 13261, Libreville, Gabon.

KEYWORDS

Floristic inventory,

Minkébé,

Gabon,

rain forest,

GilbertiodenAran.

MOTS CLÉS

Inventaire floristique,

Minkébé,

Gabon,

forêt ombrophile,

Gilbertiodendron.

ABSTRACT

This study describes the floristic diversity artd végétation of the Minkébé area

in North East Gabon, an area hitherto poorly known. A total area of 290 ha

was completely surveved for trocs > 70 cm dbh. A derailed inventory for trees

>10 cm dbh of a 3 ha plot was conducted. Many new records cottld be

added to the existing checklist. Spatial distribution of the major species is

presented as well as a preliminary végétation typology. Attention is given to

the ecological and successional status of the tree species with spécial emphasis

on Gilbertiodendron dewevrei. Comparions are ntade with data on forest

composition in Gabon from varïous authors.

RÉSUMÉ

La diversité floristique et la végétation de la région de Minkébé au Nord-Est

du Gabon, une région très peu connue jusqu'à maintenant, sont étudiées.

Une surface totale de 29U ha a été complètement explorée pour des arbres >

70 cm dbh. Un inventaire détaillé des arbres > 10 cm dbh a été également

réalisé sur une surface de 3 ha. Beaucoup d'espèces ont été ajoutées à la liste

systématique existante. La distribution spatiale des espèces dominantes est

présentée, de même qu’une typologie préliminaire de la végétation. La posi¬

tion écologique et sylvigénétique des espèces arborescentes est signalée en

accordant une attention spéciale à Gilbertiodendron dewe-vm. Des comparai¬

sons sont effectuées avec les données fournies par plusieurs autres études trai¬

tant de la composition forestière au Gabon.

ADANSONIA, sér. 3 • 1998 • 20(1)

139

van Valkenburg J.L.C.H., Ketner P. & Wilks C.M.

INTRODUCTION

This paper présents part of the results of the

floristic and végétation inventories of a multi-dis-

ciplinary study in the Minkébé area, North East

Gabon, within the Framework of the WWF pro-

gram for Gabon.

Information on the végétation of North East

Gabon is rather scarce. According to WHITE

(1983) the forests of Gabon are part of the

Guineo-Congolian régional centre ol endemism.

The forest in the North East of Gabon bclongs to

the mixed moist semi-evergreen Guineo-

Congolian forest type (WllITE 1983). Though the

prévalent végétation is moist semi-evergreen forest

of mixed composition, smatl islands of single-

dominant forest are also tound scattered throu-

ghout. The upper stratum of single-dominant

forest is uniform and dense, usually 35-45 m high

and is composcd of a single or very few species.

According to CaballÉ (1978) two végétation

types can be distinguished in eastern Gabon. The

first otte cailed “forêt dense à tendance semi-

caducifoliée'' with Pycnanthus angolensis, Penta-

clethra eervcldeana, Terminalia superba and Tri-

plocbiton scleroxylon. The second type cailed

“forêt dense humide sempervirentc” with Scypho-

cephalutn ochocoa, Pycnanthus angolensis , Penta-

clethra eetveldea.net -, Ce/tis spp ., Gilletiodendran

pierreanum and Gilbertipdendron dewevrei. The

last species forms single-dominant forest islands.

The quantitative importance of Burseraceae,

Irvingiaceae, and Olacaceae is far less than in the

west of the country (e.g. absence of okoumé,

Aucournea klaineana ), whereas the importance of

Mimosaceae and Papilionaceae increases towards

the east of Gabon.

In 1966 a forest inventory was carried out in

the Bélinga mountains 50 km sourheast of the

Minkébé area (AUBREVILLE 1967; HALLE et al

1967). REITSMA (1988) gives a detailed quantita¬

tive ecologital inventory of four one-hectare

plots of undisturbed lowland forest in Gabon.

One of his plots is located at Ekobakoba, south

of Bélinga and southeast of Makoukou (Fig. 1).

Floristically North East Gabon has been descri-

bed in 6 checklists (Halle 1964, 1965; Halle

& Le Thomas 1967, 1970; Hladik & Halle

1973; Florence & Hladik 1980).

METHODS

Fieldwork

The basecamp was situated on a high bank of

the Sing river, at 1°30’N, 12°48’E. A line survey

was conducted with an orientation of 80 grads

and 280 grads (Fig. 1). Within the transect two

surveys were carricd out, onc of trees > 70 cm

dbh (diameter at breast height measured at 13m

above ground or immediatelv above the but-

tresses if these extend beyond 1.3 m) and a

second coneerning trees >10 cm dbh. The field¬

work was carried out in 1990.

The survey of rrees > 70 cm dbh comprised an

area with a total length of 58 km and was 50 m

widc (surface area 290 ha), divided into 500 X

50 m sections. The location of every tree >

70 cm dbh was mapped, the diameter measured

and if ncccesary a voucher specimen was collec¬

ter! for identification.

The survey of trees > 10 cm dbh comprised a

transect of 6 km length and was 5 m wide (surfa¬

ce area 3 ha), divided in 100 X 5 m sections. The

locality of every tree > 10 cm dbh was mapped,

the diameter measured and if neccesary a vou¬

cher specimen was collected for identification. In

addition also ail plants, including herbs and

shrubs, bearing flowers and/or fruits présent on

the transect were collected to get an impression

of the floristic diversity. An additional floristic

survey was carried out along rivets and streams.

Data Processing

The voucher specimens and a complété set of

ail fertile herbarium specimens were sent to

Wageningeu (Herbarium Vadense) for identifica¬

tion. The first set of fertile herbarium specimens

is présent at the Herbier National CENAREST,

Libreville, A complété listing of the species can

be found in VAN VALKENBURG (1990) and in

Steel (1992).

For ail species found a diameter class distribu¬

tion was made and the location of main species

along the transect drawn on scale.

In order to ascribe an importance value to a

family or species the relative frequency of that

Family or species was determined by the follo-

wing formula.

140

ADANSONIA, sér, 3 • 1998 • 20(1)

Floristic inventory in the Minkébé région (NE Gabon)

nurnber of individuals

of a taxon

Relative frequency = - X 100%

total nuniber

of individuals

The quantitative data gathered in the végéta¬

tion survey were used for a preliminary classifica¬

tion of the forest végétation. For the végétation

analysis each section vvas treated as a relevé. Not

only the absence or présence of species was taken

into account but aJso the abundance. The num-

ber of individuals was transfered into the follo-

wing classes: absent = —, 1-2 trees = t, 3-4 trees =

2, 5-6 trees ^ 3, 7-8 ttees = 4, > 8 trees = 5.

These data were processed using a computer

cluster programme TWINSPAN (HlLL 1979)

and adapted by the former Department of

Végétation Science, Plant Ecology and Weed

Science. Twinspan is a polythetic divisive cluster

programme. The polythetic method means that

divisions are made based upon ail species présent

in the relevés, and the abundance of species is

also taken into accounr. The resulting clusters

were analysed and interpreted belore well defined

végétation types could be disttnguished. With

the computer programme Clutab, a synoptic

table was prepared, giving frequencies on a scale

of 5 of each species per cluster.

Afrer a rypology of the végétation of the 500 m

as well as the 100 ni plots was made, the large

and the small plots were combined and compa-

red. Using field observations, the data on diame-

ter distribution of the various tree species,

Fig. 1.—Map of Gabon with the location of the various research sites (adapted from Reitsma 1988).

ADANSONIA, sér. 3 • 1998 • 20(1)

141

van Valkenburg J.L.C.H., Ketner P. & Wilks C.M.

diameter distribution within plots, scarcc data

from literature about ecological requirements of

certain tree species (e.g., CABALLÉ 1978; GÉRARD

1960; Saint Aubin 1963; Whitr &c Abernathy

1996) and their status in the successional process,

insight in the forest dynamics was attempted.

RESULTS

Floristic diversity

The floristic survey of the two transects in the

Minkébé area, including riversides and a general

collecting offlowering and/or fruiting plants,

resulted in 655 taxa identifîed ro species level in

89 families (van Valkenburci 1990; Steel

1992). Many specimens could only be identifled

to genus or family level; 214 new taxa could be

added to the existing checklist for North East

Gabon (Hallé 1964, 1965; Hallé & Le

Thomas 1967, 1970; Hladik & Haï té 1973;

Florence & Hladik 1980). An additional col¬

lecting trip in December 1990 has added another

3 species to dus list of new records, although still

a considérable number of specimens has to be

identifled (Appendix 1).

The survey of trees > 70 cm dbh (totalling 290

ha) comprised a total of 1148 individuals of 113

species in 34 families. The three most important

families are Caesalpiniaceae (27,1%), Mimo-

saceae (20,9%), Burseraceae (7%).—Fig. 2. The

ten most common species are Dacryodes

buettneri, Distemonanthns benthamianus,

Erythrophlcurn ivorense , Gilbertiodmdron deive-

vret , Mo >i o pet a la n t/rus pellegrimi , Cyticodiscus

gabunemii, Pentaclethra eetveldeana ,

Piptadeniaslrum africanum , Pycnanthus

angolensis , and Pterocarpus soyauxii , followed by

Alstnnia bonne!, Terminalia superba , and

Petersiantbus maaocarpm.

The survey of trees > 10 cm dbh (totalling 3

ha) comprised a total of 1155 individuals of 202

species in 45 families. The three most important

families are Caesalpiniaceae (28.7%),

Meliaceae

Mimosaceae 20.9%

Burseraceae 7.0%

Caesalpiniaceae 27.1%

Combretaceae 3.1%

Irvingiaceae 3.5%

olliers 23 6%

Myristicaceae 3.9%

Papilionaceae 5.3%

Sapotaceae 3.0%

Total number of families: 34

Fig. 2.—Family importance (as percentage of total individuals) of trees > 70 cm dbh in a line survey of 290 ha in the Minkébé area.

142

ADANSONIA, sér. 3 • 1998 • 20(1)

Floristic inventory in the Minkébé région (NE Gabon)

Euphorbiaccae (9%). Burscraceae (7%).—Fig. 3.

The ten most commun species of medium or

small size are Polyalthia sudveolens, Santiria tri¬

mera, Cryptosepalurn congolanurn, Plagiosiphon

multijugus, Scorudophloeus zenkeri, Centroplaeus

glaucinus, Dickostemnid glauecstens, Plagiostyles

africana , Coula edulis , and Cola rostmta.

From the rret inventory it is clear chat some

species do not occur as big trees with dbh >

70 cm in the 500 ni relevé, but are restricted to

the 100 m plots as smaller individuals. One

example is Gilbertiodendron ogoouense. This spe¬

cies can be considered as undersrorey species,

never reaching great height or diameter. These

smaller trees provide a means of subdividing the

végétation types defined by rhe larger trees.

Various other species may be considered as typi-

cal understorey species, such as Cola rostrata,

Coula edulis and, Santiria trimera. Thercfore the

species composition of the 10 cm diameter plots

is not représentative of the upper canopy.

Species which occur in ail diameter classes such

as Gilbertiodendron dewevrei, Santiria trimera

and. Cola rostrata can be considered as primary

species.

The diameter distribution and spatial distribu¬

tion of these common species (Fig. 4) illustrate

their différences in stature and eeological préfé¬

rences. The potenual srature of the species

decreases from Gilbertiodendron dewevrei to Cola

rostrata. Whereas Gilbertiodendron dewevrei and

Cola rostrata arc muiually exclusive. Santiria tri¬

mera is found in association with both species.

Eeological préférences can be discerned in the

survey of large diameter trees (Fig. 5).

Pterocarpus soyauxii and Cylicodiseus gabunesis are

common species, the latter found in slightly

higher densities. Gilbertiodendron dewevrei and

Monopetalanlhus pellegrinii hâve a very well defi-

ned distribution and are mutually exclusive.

Dacryodes buettneri bas a rather local distribution

but can be found in high densities. Terminalia

superba, a good indicator for late secondary forest,

also occurs very locally with high densities.

Caesalpiniaceae 2° rn.

Euphorbiaceae 9.0%

Mimosaceae 3.3%

Olacaceae 7.6%

Papilionaceae 4,3%

Rubiaceae 2.7%

Sterculiaceae 7.1%

Annonaceae 6.0%

0 [tiers 23.4%

Total number of families: 45

Fig. 3.—Family importance (as percentage of total individuals) of trees > 10 cm dbh in a line survey of 3 ha in the Minkébé area.

ADANSONIA, sér. 3 • 1998 • 20(1)

143

van Valkenburg J.L.C.H., Ketner P. & Wilks C.M.

Diameter distribution

Gilbertiodendron dewevrei

Spatial distribution

Gilbertiodendron dewevrei

Diameter class (cm)

Plot number

Santiria trimera

Diameter class (cm)

Plot number

Diameter class (cm) Plot number

Fig. 4.—Diameter distribution and spatial distribution of three common tree species (per 100 x 5 m plot) along the 6 km transect of

trees > 10 cm dbh: Gilbertiodendron dewevrei, Santiria trimera, and Cola rostrata. (y-axis: number of trees).

144

ADANSONIA, sèr 3 • 1998 • 20(1)

Floristic inventory in the Minkébé région (NE Gabon)

Pterocarpus soyauxii Cylicodiscus gabunensis

Gilbertiodendron dewevrei Dacryodes buettneri

Monopetalanthus pellegrinii Terminalia superba

Fig. 5.—Spatial distribution of some characteristic tree species (per 500 x 50 plot) along the 58 km transect of trees > 70 cm dbh:

Pterocarpus soyauxii, Cylicodiscus gabunensis, Gilbertiodendron dewevrei, Dacryodes buettneri, Monopetalanthus pellegrinii, and

Terminalia superba. (x-axis: plot number; y-axis: number ot trees).

ADANSONIA; sér. 3 • 1998 • 20(1)

145

van Valkenburg J.L.C.H., Ketner P. & Wïlks C.M.

Description of the végétation types

The name of each végétation rype is based on

two species of which 011 e is a more commun with

highest frequency class, and the second is either a

différenciai or an accompanying species. A diffé¬

renciai species is différenciai for a cluster if ics fre-

quency is at least 30% (= one frequency class)

higher than in any other cluster, or two related

clusters (Sc.HAMlNFF. et al. 1995). Occasionally a

species was considered diffcreiuial when it occur-

red in a frequency of 20% higher than in any

other cluster.

When no differential species were présent or no

clear dominant species the second ntost impor¬

tant species was taken co name the type.

Typology of the 500 m length plots

In total 115 plots of 500 X 50 m were sampled

(one plot was empty). The number of relevés

used for clustering is 101 as ail the relevés with

fewer than 5 individual trees of > 70 cm dbh

(n = 14) were not included.

Front the ordered Twinspan table five main

clusters can be distinguished. These can be descri-

bed in terrns of frequency and relative abundance

of the species présent and treated as végétation

types. Table 5 présents the synoptic table of the

discerned clusters and végétation types.

Type A: Gilbertiodmdron dewevrei - Pterocarpus

soyauxii type (11 relevés).

This type is very characteristic because ol the

high frequency ol Gilhertiodcndm n dewevrei with

high average abundance. In fact Gilbertiodendron

is the only characteristic species. Oxystigma buch-

holzii occurs in this type only, but with low fre¬

quency. In general this type is poor in species

(average 5.2). In comparison with ail the follo-

wing types Cylicodiscus gabunensis and

Erythrophleum ivoreme are virtually absent.

Type B: Dacryodes buettneri - Baillonella toxî-

sperma type (21 relevés).

This type is characierised by the high occurrence

of Dacryodes buettneri and relatively high présen¬

ce of Buillonella toxisperma (highest frequency

class ol ail types), both differential species.

Common species with relatively high frequency

are Cylicodiscus gabunensis, Piptadeniastrum afri-

canum and Erythrophleum ivorense. Some species

occur more in this type than the odiers, such as

Eagara heitzii, and Testulea gabonensis. Parkia

bicolor occurs slightly more frequently than in A

and D.

Type C : Monopetalanthus pellegrinii -

Cylicodiscus gabunensis type (27 relevés).

This type is less clearly defined than the latter

and the f’ollowing type. Monopetalanthus pellegri¬

nii is the differential species of this type and is

abundanr. Monopetalanthus letestui occurs in this

type with highest frequency and is slightly diffe¬

rential.

Type D: Cylicodiscus gabunensis - Pycnanthus

angolensis type (34 relevés).

This type has no differential species, but

Pycnanthus angolensis as well as Distemonanthes

benthamianus are almosr confined to this type

and the next. Cylicodiscus gabunensis occurs with

high frequency, but Monopetalanthus is absent.

Gambeya lacourtiana and Uapaca paludosa are

présent mainly in this type. Average number of

species is higher than in the previous types.

Type E: Terminalia superba - Ceiba pentandra

type (8 relevés).

This rype is characterised by four differential spe¬

cies, three with high frequencies (highest of ail

types) and relatively high abundance i.e.

Terminalia superba , Alstonid boone't and Ceiba

pentandra , and Ceins adolft-friderici, The com¬

mon species which occut in A, B. C and D are

présent in this type, but ail with low frequency.

Pycnanthus angolensis is présent with higher fre¬

quency, but with slightly lower average abundan¬

ce than in type D. Blighia welwitschii and

Mitragyna stipulosa are confined to this type, and

may also be considered as differential.

Typology of the 100 m length plots

In total, 59 plots of 100 X 5 m were sampled

(one plot was empty). Front the ordered

Twinspan table 4 végétation types could be dis¬

tinguished, two ol which hâve .subtypes. The fre¬

quencies of each species in the relevés of each

(sub)type are given in Table 6.

146

ADANSONIA, sér. 3 • 1998 • 20(1)

Floristic inventory in the Minkébé région (NE Gabon)

Type 1 (cluster 1+2): Gilbertiodendron dewe-

vrei - Gilbertiodendron ogoottense maintype (22

relevés).

This type is characterised by the high frequency

as well as abundance of Gilbertiodendron dewe-

vrei and relarively high frequency (40-60%) of

Gilbertiodendron ogootiense , boch différenciai spc-

cies for clusters 1 and 2. This cype can be dîvided

into t\vo subtypes namely:

l.a: subtype wilh Oxystigrna buchbolzii and

Cryptosepalum congolannrn (12 relevés).

Oxystigrna buchbolzii, Cryptosepalum Congolanurn,

Treculia africana, Plagiosiphon Mültijugus and

Uapaca beudelotii are différenciai speties for this

type, compared wilh ail othcr types. There are

hardly any commun spedcs in this subtype,

l.b: subtype with Santiria trimera and

Lasiodiscus marmoratus ( 10 relevés).

Lasiodiscus marmoratus is differential for this sub¬

type and Santiria trimera is frequent, both are

absent in the previous one. Typical species are

Heisteria paroiflora , Strombosia scheffleri , Baphia

spp. On average this subtype has a higher num-

ber of species than subtype 1 .a (13.0 vs. 9.5), but

ail with low fréquences.

Type 2 (cluster 3): Calpocalyx dinklagei -

Gretvia coriacea cype (4 relevés),

This type is based on only four relevés and it is

therefore doubtful whethcr it can be considered

as a separate type, although it has five differential

species: Calpocalyx dinklagei, Grewia coriacea as

well as Xylopia quintasii, Anthonotba macropbylla

and A. cf. ferruginea. A large number of species is

présent only in this type, but ail with low fre-

quencies (21-40%), In contrast to the next types

the common species Cola rostrata , Petersiantbus

macrocarpus, Plagiostyles africana and Strombosia

pustülata do not occur.

Type 3 (cluster 4 + 5): Cola rostrata -

Petersiantbus macrocarpus maintype (21 relevés).

This type is characteristed by Lts high frequency

of Cola rostrata. Scorodophloeus zenkeri also

occurs very frequencly, as well as Strombosia pus-

tulata and Petersiantbus macrocarpus. This type

can be divided into rwo subtypes:

3.a: subtype with Coula edulis (10 relevés).

In this subtype Polyalthia suaveolens and Coula

edulis are very frequent. Centroplacus glaucinus is

absent contrary to subtype 3. b.

3.b: subtype with Strombosia pustuiata (Il rele¬

vés).

Strombosia pustuiata and Plagiostyles africana

occur more frequent and abundant in this type

than in subtype 3.a. Dichostemma glaucescens can

locally be abundant (4 plots). In this subtype

Polyalthia suaveolens and Coula edulis are poorly

represented in comparison wirh subtype 3.a.

Type 4 (cluster 6): Santiria trimera -

Centroplacus glaucinus type (12 relevés).

This type is characterised by the presence of

Centroplacus glaucinus and Scorodophloeus zenkeri

with highest frequencies of ail types. There are

no differential species for this type. Polyalthia

suaveolens and Plagiostyles africana occur also fre-

quently. Cola rostrata and Petersiantbus macrocar-

Capital letters = 500 m plots végétation types

Numbers =, 100 m plots végétation types

j = bounderies between 500 m plots

] = assumed bounderies of the main végétation types

■ = creek

A | c | D | D | A I 0 | D I C I A | ! | A | D | E |

la la la lb 4 3b 4 4 3b 3a 2 2 3a tb 4 e, e, 4 4 1b lb Ib 1b 1b 4 4 3a 3b 3a 3b 3b 3b 3b 3‘a 3a 3b 3b 3a « la la la la la la la la la 1b ib ib 1b 1b 4 3b 3b 3a 4 3a 2 2

Fig. 6.—A schematic présentation of the occurrence of the 500 m and 100 m végétation types in the Minkébé survey.

ADANSONIA, séf. 3 * 1998 • 20(1)

147

van Valkenburg J.L.C.H., Ketner P. & Wilks C.M.

Gilbertiodendron dewevrei

10 I-

1 11 21 31 41 51

Gilbertiodendron ogoouense

Piagiosiphon muitijugus

Oxystigma buchholzii

Fig. 7.—Spatial distribution of sonne characteristic tree species (per 100 x 5 m plot) along the 6 km transect of trees > 10 cm dbh:

Gilbertiodendron dewevrei. Gilbertiodendron ogoouense, Cryptosepalum congolanum, Piagiosiphon muitijugus, Santiria trimera, and

Oxystigma buchholzii. (x-axis: plot number; y-axis: number of trees).

148

ADANSONIA, sér. 3 • 1998 • 20(1)

Floristic inventory in the Minkébé région (NE Gabon)

pus are poorly represented, compared to type 3.a

and 3.b, with which type 4 shows great similarity.

Combination of 500 m plots with the 100 m

plots

Figure 6 gives in schematic form the occurrence

of the 500 m and 100 m types in the transect.

Combining the types derived from the small

plots with those from the large plots we see sonie

striking features.

Type l.a and l.b coïncide with main type A.

This type now' can clearly be subdivided into a

Gilbertiodendron dewevrei type with Oxystigma

buchhoLzii and a Gilbertiodendron type without

Oxistygma, whereby rhe first type occurs close to

the streams and the latter on somewhat drier

places adjacent.

The ecological préférences of some smaller spe-

cies occurring in the two subtypes of the

Gilbertiodendron maintype are illustrated in

Figure 7. Gilbertiodendron dewevrei and G.

ogoouetvse are found in both wet and dry habitats,

Cryptosepalum eongolnnum and Plagiosiphon rnul-

tijugus are only présent in the wct habitat and

Oxystigma bttehholzii is çonfinçd to the perma-

nently inundated sites along the river. Contrary

to this Santirict trimera is only présent in the

drier land habitat.

Type A is clearly primary forest; of the constitu-

ting species ail diameter classes are represented,

forming large almost monospecifïc patches.

There are no smaller plot types which coincidc

with type B, but Iront the large trce inventory

alone type B can be classified as primary forest,

because of absence of clear secondary species.

Type C and type D ate closcly related types.

The main différence is that D shows more signs

of former disturbances through the occurrence,

in higher frequencies and/or higher abundance

of a number of secondary species, such as e.g.

Pycnanthus angolensis, Disternonanthus bentha-

miatms, Petersianthus macrocarpus. The abundan¬

ce of Cylicodiscus, one of the main primary

species in C and D, is almost equal.

This différence is also reflected in the types 3.a,

3.b as well as type 4, which are also very related

to each other, ail three corresponding with types

C and D. Types 3.a, 3.b and 4 represent develop¬

ment stages of forest growth from secondary to

primary, type 4 having already more primary

species with higher abundance than 3.a; while in

type 3,a the primary species occur in higher fre-

quency and slightly more abundant than in 3.b.

There arc no smaller plot types which coincide

with type B, but from the large tree inventory

alone type B can be classified as primary forest,

because of absence ot clear secondary species.

Type 2 ol which only 4 relevés are présent

sccms to coincide mainly with F. (3 out of 4 rele¬

vés) and slightly with D. The large size

Terminalia superba, Ceiba pentandra and

Eribroma oblongum indicate that type E may be

classified as late secondary forest. The ptesence

ot Anlhonotba macrophylla and Xylopia quintasii

in the 100 m plots indicates that the forest is

changing into primary forest.

As there probablv are more changes taking

place in the composition of smaller trees than in

tire large trees it is difficult to show very strict

corrélations between the types of the 100 m plots

and the 500 m [îlots.

DISCUSSION

The data on family and species importance of

trees > 70 cm dbh can be compared with the

data of CABALLÉ (1978) concerning trees >

60 cm dbh (see Table 1 + Table 2).

Comparlng the data on family importance of

large trees from CABALLll (1978) with the

Minkébé results, the importance of

Caesalpi niaceae is (almost) the same,

Mimosaceae are more important in the Minkébé

survey, but Myristicaceae importance is much

lower (Table 1 ). If we compare the importance of

the tree species (Table 2), the absence of

Scyphoeephalutn ochocoa in the Minkébé survey is

noticeable and this explains also the low impor¬

tance value of Myristicaceae. T he higher value

for PiptadeniaStrum afrieanum and Cylicodiscus

gabunensis in Minkébé accounts for rhe higher

importance value of Mimosaceae. Though the

relative frequency of Caesalpiniaceae is the same,

the share of the constituting species differs.

Gilletiodendron pierreanum is absent in the

Minkébé survey, and Disternonanthus benthamia-

ADANSONIA, sér. 3 • 1998 • 20(1)

149

van Valkenburg J.L.C.H., Ketner P. &c Wilks C.M.

Table 1.—Comparison of family importance of large

trees in the Minkébé area (290 ha) with the results of

Caballé (1978) for East Gabon. Values in % of total

number of trees

Caballé *

dbh > 60 cm*’

Minkébé

dbh > 70 cm

Caesalpiniaceae

26.7

27.1

Burseraceae

4.2

7.0

Irvingiaceae

2.6

3.5

Olacaceae

2.4

Mimosaceae

14.6

20.9

Papilionaceae

3.1

5.3

Myristicaceae

8.5

3.9

* : Based on the 15 most abundant species.

*': Based on ail individuels of ail species présent.

nus, Erythrophleum ivorense and Monop étalantbus

pellegrinii are not among the 15 most important

species of Cahai.I.Ê (1978).

The data on family and species importance of

trees > 10 cm dbh can be compared with the

data of CA8AU.fl (1978) concerning trees 20 cm

< dbh < 60 cm (sec Table 3 + Table 4).

The family importance value of small and

medium sized trees (Table 3) shows an extremely

high value for Caesalpiniaceae in the Minkébé

survey (when compared ta CABAI1.Ê). This is

causcd by the lact thar 23 of the 59 relevés

(39%) are located in Gilbertiodendron forest,

whereas in the survey of trees > 70 cm dhh onlv

11 of the 104 relevés (11%) are located in

Gilbertiodendron forest. Comparing the relative

frequency values of CABALLE (1978) with the

Minkébé data, the différence in importance value

of Caesalpiniaceae is illustrated with the higher

value of Gilbertiodendron spp. and the 5 additio-

nal Caesalpiniaceae species in Minkébé (Table 4).

The data of the survey of trees > 10 cm dbh can

also be compared with the results of Rbit.SMA at

Ekobakoba (REITSMA 1988), which is most

proximate to the Minkébé area. REITSMAs one-

hectare plot, on the top of a small hill with little

variation in relief, comprised 438 individuals >

10 cm dbh, 85 species in 30 families. The chree

most important families are also Caesalpiniaceae,

Euphorbiaceae and Burseraceae.

According to REITSMA (1988, table 9, page 46),

who uses importance values, the ten most impor¬

tant species are Hymenostegia pellegrinii,

Scyphocephalum ochocoa , Santiria trimera,

Di ali ttm pachyphyllwn , Plagiostyles africana,

Desbordesia gbiucesccns, Dacryodes buettneri. Cola

rostrafa, Polyalthia uiaveolens, and

Entandrophragnui candollei (a single tree only).

The average density of trees > 10 cm dbh in the

Minkébé area is 385 trees per ha, which is less

than in RtlTSMA's Ekobakoba site, who found

438 individuals in one hectare. This can be attri-

bttted to the fact tliat part of the survey was

situated in the dynamic river System of the river

Sing. The higher number of species found can-

not only be attributed to the larger arça surveyed,

but is surely also caused by the greater environ-

mental tliversity, since swamps, periodically Hoo-

ded forest and dry forest are included in the

survey.

The high value for Hymenostegia pellegrinii in

the Ekobakoba plot and its absence in the

Minkébé survey of trees > 10 cm is anorher

example of local dominance of certain

Caesalpiniaceae species. The high value is also

caused by the small sample size/area surveyed.

REITSMA ’s plot is a 100 x 100 m plot with little

Table 2.—Comparison of species importance of large

trees for the 15 highest ranking species in the Minkébé

area (290 ha) with the results of Caballé (1978) for East

Gabon. Values in % of total number of trees.

Caballé

dbh > 60 cm

Minkébé

dbh > 70 cm

Scyphocephalum ochocoa

5.2

Pentaclethra eetveldeana

4.5

3.7

Gilleliodendron pierreanum

4.2

Pycnanlhus angolensis

3.3

3.7

Petersianthus macrocarpus

3.1

2.7

Alslûnia baonei

3.0

2.6

Dacryodos huohneri

2.9

6.5

Piptadamaatrum africanum

2.9

4.2

Cylicodiscus gabunensis

2.6

7.9

Cnips briayi

2.5

Qllberliûdendron sp.

2.4

5.6

Pleracarputi soyauxii

2.3

4.8

Uapaca sp

2.3

1.1

Scorodûphloeus zenkeri

2.0

1.1

Tcrminalia superba

2.0

2.5

Üistemonanthus benthamianus

3.5

Erythrophleum ivorense

5.3

Monopetalanthus pellegrinii

2.3

150

ADANSONIA, sér. 3 • 1998 • 20(1)

Floristic inventory in the Minkébé région (NE Gabon)

Table 3. —Comparison of family importance of small and

medium sized trees in the Minkébé area (3 ha) with the

results of Caballé (1978) for East Gabon. Values in %

of total number of trees.

Caballé *

20 cm < dbh < 60 cm

Minkébé **

dbh > 10 cm

Euphorbiaceae

8.1

9.0

Caesalpiniaceae

9.3

28.7

Burseraceae

6.9

7.9

Mimosaceae

5.0

3.3

Annonaceae

3.3

6.0

Lecythidaceae

3.1

Ulmaceae

2.3

Olacaceae

2.0

7.6

Myristicaceae

2.8

Pandaceae

1.2

* : Based on the 15 most abundant species.

**: Based on ail individuals of ail species présent.

variation in relief. The Minkébé survey is a three

ha plot of 6000 X 5 m with much more variation

in topography and abiotic factors. The chances

for Local dominance over the entire plot are

much smailer. CaBAUT’s surface area covers vast

stretches of the eastern zone of Gabon and there-

fore high species importance values are impos¬

sible.

Whcn comparing family importance values of

trees > 60 cm dbh with 20 cm < trees < 60 cm of

Caballé (1978) it is clear rhar Caesalpiniaceae

and Mimosacene are very important canopy trees

(41.3%) but are far less prominent (14.4%)

among small and medium sized trees (the floris¬

tic composition of small and medium sized trees

is more diverse).

The recognized végétation types of the 500 m

as well as 100 m plots show the internai dyna-

Table 4.—Comparison of species importance of small and medium sized trees for the 15 highest ranking species in

the Minkébé area (3 ha) with the results of Caballé (1978) for East Gabon and Reitsma (1988) at Ekobakoba (1 ha).

Values in % of total number of trees.

Reitsma *

dbh > 10 cm

Minkébé

dbh > 10 cm

Caballé

20 cm < dbh < 60 cm

Plagiostyles africana

5.3

1.8

8.1

Scorodophloeus zenkeri

5.0

6.4

Santiha trimera

5.3

7.2

5.8

Pentaclelhra eetvetdeana

1.4

3.9

Polyalthia suaveotens

3.3

2.9

3.2

Petersianthus macrocarpus

1.6

3.1

Celtrs brieyi

2.3

Coûta edutis

3.5

2.0

Scyphoœphalum ochocoa

3.3

1.4

Dialium pachyphyllum

4.6

1.7

Coelocaryon preusii

1.4

Gitbediodeodron dewevrei

8.2

1.3

Gitbertiodendron ogoouense

1.7

Panda oleosa

1.2

Pentaclettira macrophylla

1.1

Dacryodes buettneri

2.0

1.1

Desbordesia glaucescens

2.5

Entandrophragma candollei

0.3

Hymenostegia pellegrinii

7.6

Cola ros trata

4.6

5.4

Cenlroplacus glaucinus

1.9

Cryptosepalum congolanum

3.0

Dichosiemma glaucescens

2.3

Oxystigma buchholzii

1.6

Plagiosipbpn multijugus

2.0

*: 10 most important species according to Reitsma’s importance value.

ADANSONIA. sér. 3 • 1998 • 20(1)

151

van Valkenburg J.L.C.H., Ketner P. & Wilks C.M.

Table 5.—Synoptic table of the végétation types of the 500 m plots (trees dbh > 70 cm) of the Minkébé area (for

codes see below).

Cluster

Végétation type

Number of relevés

Aver. no. of species

5.2

7.9

7.8

9.0

7.9

Standard déviation

2.7

3.2

2.4

2.9

3.3

Differential species

ûilbertiodendron dewevrei

Oxystigma buchholzii

Baillonella toxisperma

III

Monopetalanthus letestul

Monopetalanthus pellegrinii

III

Dacryodes buettneri

Alsionia boonei

Pycnanthus angolensis

III

Terminalia superba

Distemonanthus benthamianus

III

Gambeya lacourtiana

Celtis adolli-lriderici

Xylopia hypolampra

Ceiba pentandra

III

Blighia welwitschii

Mit.ragyna stipulosa

Common species

Petersianlhus macrocarpus

Pentaclethra eetveldeana

III

Pterocarpus soyauxii

III

Erythrophleum ivorensis

III

Cylicodiscus gabunensis

III

Hymenostegia pellegrinii

Piptadeniastrum africanum

III

Irvingia grandi folia

Guibourtia tessmannii

Parkia bicolor

Fagara heitzii

Scorodophloeus zenkeri

Millettia laurenlii

Klalnedoxa gabonensis

Pentaclethra macrophylla

Celtis tessmannii

Other species (excluding rare)

Amphimas pterocarpoides

Beilschrmedia sp.

Uapaca patudosa

Detarium macrocarpum

Anopyxis klameana

Celtis mildbraedii

Canarium schweinfurtii

Coula edulis

Dialium pachyphyllum

Dialium dinklagei

Erismadelphus exsul

Fillaeopsis discophora

Entandrophragma cylindricum

Entandrophragma candollei

Parkia llllcoidea

Irvingia gabonensis

152

ADANSONIA, sér. 3 • 1998 • 20(1)

Floristic inventory in the Minkébé région (NE Gabon)

Milicia excelsa - - + + I

Lophira alata I - - - I

Syzygium sp. - I - I I

Dracaena mannii - + - I I

Nauclea diderichii - + I I -

Pterygota bequaertii I - I

Tessmannia afrlcana I + - +

Tessmannia anomala I - I

Tetraberiinia bifoliolata I - + I -

Turraeanthus africanus I - - I

Rare species

Afzelia bipindensis: D.+; Albizia adianthifolia : B.+; D,l; Albizia ferruginea: D.l: Antiaris africa-

na: B Antrocaryon micraster. B,l; C.+, Bombay bunoponense : A,l; D.l; Coelocaryon preus

sir: D,+: Crudia gabonensis• B,+: D,+; Dacryodes igaganga: C.+. DdrfelVa oblongum: C,+;

Desbordesia glaucescens: B.l; Diogla zenkeri : C.+; Drypetes goswgilleri: C,+;

Enlandropbragma congoense: C,l; Eribroma oblongum : C,+ ; D,l, Ficus elaslicoides: D,+;

Gambeya beguei : C,+; D.+. Guarea Ihomsonii: D.l: Gilbertiodnudron ugoouense : B,+;

Heisteria parvifolia: E,l; Julbemardia s eretll: C.*; Kayodendron bndelloides : D.+ : Flainedoxa

trillesu: C,+, D,l; Landolphia forolii: C t >; Lovaa trichiloidos : A,l; B.l; Macatanga sp.: A,l

Mammea afrlcana: A,l; Maranthes glabra: B.l; Maranthes sp ’ B.+; C.+: Myrianthus arboreus:

E.l: Newtonia glartdulifera : C, + : Nowlania ni. glanduliferà. B.+. Oldtieldla atricana: B.+; C.

Ongokea gore. B.l; C,l; Parmar! excelsa: B.+; C,+; Pachyelasma lessmannii : B.+; Panda

oleosa: C,*: D.l; Pteleopsis hylodondmn: C.+; 0,1; Plerygopodium oxyphyllum: B.l; D,+;

Rhodognaphalon brevicuspe: C.+ : Scottollia kininei: D.l: Staudtia gabonensis: D.l;

Stemenocoleus micranlhus: B.l; Strombosta scbettlen : C,+; Testufea gabonensis: B.l; C,+;

Tieghemella atricana: B,+; Trichilia lessmannii. C, I, Trichoscypha acuminata: B,+;

Tridesmostemon omphalocarpoides: C.+; Uapaca heudelotii: B.+.

Frequencies in 5 classes

III

= species not présent in the relevés of the cluster

= species présent in 15% ot the relevés

= species présent in 6-20% ot the relevés

= species présent in 21 -40% ot the relevés

= species présent in 41-60% ot the relevés

= species présent in 61-80% ol the relevés

= species présent in 81-100% ot the relevés

mies of the forest, with young and old stages

alternating over short and long distances (Fig. 6).

As the inventory took place along a rransect with

adjacent plots it could be expected th.it the typo-

logy would be less pronounced. Figure 6 shows

the borders between each sample plot On the

basis of the typology the assumed boundaries

between the different végétation types are drawn.

More data are needed belote something can be

said about the succcssional status of the different

types (Type D either a transitory type liaving élé¬

ments of Type A or transitory to E, having

various species in common with E). It also

contains fewer individuals and fewer species in

compari.son with the other types. It is either not

yet completely developed to mature forest or in a

degraded phase.

As little was recorded about the soil and the

local topography. not much can be said about

the ecology of the végétation types, except for the

Gilbertiodendron dewevrei - Oxysngmd subtype,

which occurs primarily along streams and is

inundated. Medium size trees are Oxystirm buch-

holzii and Plagia siphon muLtijngus. The

Gilbertiodendron dewevrei - Sd/niria trimera

forest, on drier places, has a canopy dominated

by Gilbertiodendrari dewevrei, but Oxystigma is

absent, while medium trees indude Sanrrria tri¬

mera, Strombosia scheffleri , Coula edulis and

Cardpa procera.

The végétation types described in this study are

but local types of which the syntaxonomic signi-

ficance can only be determined if far more rele¬

vés front nearby régions beconte available.

Ecological parameters such as soil characteristics,

hydrology, local topography and human influen-

ADANSONIA. sér. 3 • 1998 • 20(1)

153

van Valkenburg J.L.C.H., Ketner P. & Wilks C.M.

Table 6. —Synoptic table of the végétation types of the 100 m plots (trees > 10 cm) of the Minkébé area (for codes

see below).

Cluster

Végétation type

Number ol relevés

Aver no of species

9.5

13.1

12.8

14.0

12.5

13.8

Standard déviation

3.1

2.4

3.2

4.0

1.3

2.8

Differential species

Cryptosepalum congolanum

Oxystigma buchholzü

Uapaca heudelotii

Treculia africana

III

Plagiosiphon multijugus

Lasiodiscus rnarmoratus

Gilbertiodendron ogoouensis

III

Gilbertlodendron dewevrei

Anthonotha cl. lerruginea

III

Anthonotha macrophylla

III

Xylopia quintasii

Calpocalyx dinklagii

Grewia coriacea

Cola roslrata

Petersianthus macrocarpus

III

Trichoscypha acummata

Common species

Santiria trimera

III

Coula edulis

Polyalthea suaveolens

III

Pentaclethra eetveldeana

III

Dialium pachyphyllum

Beilr.chmiedia sp.

Scorodopbloeus zenkeri

Plagioslyles alricana

III

Centroplacus glaucinus

III

Strombosia pustulata

III

Celtis tessmannii

Pausinystalia macroceras

III

Dichosiemma glaucescens

Panda oleosa

III

Sorindeia nitidula

Trichllia weMtschii

Picraiima nitida

Irvingia gabonensis

Indel.

III

Othcr species (cxcluding rare)

Strombosia schettlen

Heistgria pan/ifplia

Baphia buettneri

Maroyopsis longiloha

Carapa procera

Tetràb&fltnla bitollata

Baphia puhnsœrin

Pterocarpus soyauxll

Dacryudes klainuana

Dialium dinklagai

Afroslyrax leoidophyllus

Drypotcs gosweilleti

Garcinia gnetoidas

154

ADANSONIA, sér. 3 - 1998 • 20(1)

Floristic inventory in the Minkébé région (NE Gabon)

Staudlia gabonensis

Xylopia parviflora

Pseudospondias sp.

Pachypodanthium slaudtii

Markhamia tomentosa

Xylopia hypolampra

Alstonia boonei

Rauvolfia caffra

Sapium comutum

Irvingia grandifolia

Ochthocosmus sp.

Psychotria sp.

Erythroph'eum ivorensis

Pycnanthus angolensis

Terminalia suberba

Diospyms piscatona

Maesopsis eminii

Nauclea diderichii

Rothmannia whhfleldii

Tricalysia biafrana

Grewla sp.

Celtis mildbraedii

Rare species

Albizia adianlhlfolia : 4,1; Albizia ferruginea: 4,1; Angylocatyx sp.: 1b,I; 3b,I; Artnonidium mannii: 3b,I; 4,1;

Anopyxis klaineana: 4,1; Aphanocalyx cynometroides: 1b,l; Artabotrys sp.: 3a,1; Baikiea insignis: 1b,I;

Raphia cl. leptostemma: 1b,I; Raphia sp : 1b,I; Raphia sp/ lb,l; Raphia s p: 3a,I; Raphia sp.: 1b,I, 3b,I;

Baphia sp,: 1b,I; 3a,I; Berlinia sp.: 1b,l- Blighia sp:. 3a,I Bridelia mlcrantha: 1b,I: Caloncoba welwitscfiii:

3a,I; Camptostylis mannii: la,l; 3a,I; Cellis adoih-friderici: Ib.l; Chytrantlnis sp., 3b.I. Cletodendron sp:. 4,1;

Coeiocaryon preussi: 3a,I, Coflea sp: la,!: Vola verticillala: 1b, I; Cola sp: Ib.l; Cola sp : 1b, l; Cola sp:.

1b,l; 3a,I; Cola sp: 4,1; Cola sp.: 3a,I; Combrelum homaiiûides: 4,1, Combretum sordidum: 3a,I; 3b,!;

Combretum sp: 4,1: Cylicodiscus gabunensis: 3a,I; 4.1: Cynometra sanagaensis: la,!; Dacryodes

buettneri: 4,1; Dalbergia sp: 1 a.l; 1b,l, DalhouSiea afrlcana: 3b,l; Datanum macrocarpum: ib.l; 4,1;

Dichapetalum choristllum: 3b,!; Didimosalpinx abbvokutae 1b,I; Diospyms ferrear. 1a,l; Diospyros gilletir,

1 a.l; Diospyms mannii: 1b, !; 3a, I, Discoglyprenvm caloneura: ib.l; Distemonan/hus benthamianus: 3a,I

3b,I; Drypntes sp.: 3b,I: 4.1: Entandrophiayma cyiindricum: 4,1; Enbroma obtonga: 3b,l; Ehocoelon sp:

3a,I; Gambe y a boukokoensis: 4,!; Gamboya tacourtiana: 3a,I; 3b, I; Garcinia mannii: 4,1; Garcinia slaudtii:

3a,I; Gardénia imparfaits: 1a,l; Grewia pinnatitlda: 1 a.l: 1b,i: Grewia sp. 3a.l; Griffonia sp: 4,1; Guarea

cedrata: 3a,I; Kiainudoxa gabonensis. 3a,I, Landolphia toretil: 3b,I; Lindackerla dentata: 3a, I; 4,1:

Lonchocarpus sp • la.l; Lophira alata : Ib.l: Mnr.aianga sp: la i 1 Magnistipula zenkeri: 1 a.l; Mallotus

oppositîfolius: la,!; Mammea atricana: la.l; 3a,I; Maprounea atricana: ib.l; Massularia acuminata: 3a.I:

Microdesmis sp: la.l: Milicia excelsa: 3b,I; Milletlia bipindonsis • Ib.l; Millettia laurentii 1b,I: 3b.l; MiHettia

sp.: la,!; Monopetalanthus microphyilus: 3b,I; 4,1; Napoieonaea sp.: la.l: Nauclea pobeguinii: la,h

Nauclea vanderguchtir la.l; Neuropeliis acuminata: 1b,l; Odulobus heleromerus: 3a,I; Odyenda gabo¬

nensis: 3b, I; 4,1; Omphalocarpum proceium: 3b,l; Oubanguia et atricana: la.l; Ib.l: Pachypodanthium

confine: 4.1; Pancovia sp: 3a.I; 4.1; Parinari excelsa: 1b,I, Pausmystalla johimbe: 3b,I: 4,1: Piptostigma gta-

brescens: Ib.l; Plagiosiphon emarginatus: la.l; Portemndia ctadantha: 3a,I; Pterocarpus sp:. 1a,l,

Pterygota bequaertli : 4,1; Rhaphiostytts sp.: 3a, I; Rauvoltia letouzeyi: 3b,l; Rinorea kamerunensis: 3b,I;

Rinorea subsessilis: 3b,I; Rinorea welwitscbii : 1b, I; Sorindeia sp.: 1b, I; Sorindeia sparanoi: 1 a, I ;

Strepbonema mannii: 1b,l; Strycbnos campicola : la,!; Synsepalum longec-uneaium: 1b,I; 4,1; Syzygiurn

sp.: 4.1, Tabernaemontana crassa: 3b,I; Tessmannla anomala: la.l: Testulea gabonensis: 4,1;

Thomandersia laurilolia : 4,1; Trichilia monadelpha. 4a.l, Tricboscypha sp: 3a,I; Xylopia aethiopica: 4,1;

Xylopia letestui ; 1 t>,l; 3a,l; Xylopia pinaartii: 3a, I; Xylopia sp:. 4,1; Xylopia staudtii: 3b,I.

Frequencies in 5 classes

III

= species not présent in tlie relevés of the cluster

= species présent in 1-5% of the relevés

= species présent in 6-20% of the relevés

= species présent in 21-40% ol the relevés

= species présent in 41 -60% of the relevés

= species présent in 61-80% ot the relevés

= species présent in 81-100% of the relevés

ADANSONIA, sér. 3 ■ 1998 • 20(1)

155

van Valkenburg J.L.C.H., Ketner P. & Wilks C.M.

ce are lacking. These vvill be necessary ro confirm

the existence of sucli types and make it possible

to interpret the forest types in an ecological

sense. Doing so the types can possibly bc used

for rnuch wider areas. Through species composi¬

tion these types can only be distinguished as pri-

mary or secondary types.

Although not a dassical way of nuking végéta¬

tion relevés and ty'pology, whereby for eaclt spe¬

cies in a homogcnous sample plot a

cover/abundance code is given (Kf.NT & CoKER

f992), recording presence/absence plus relative

abundance according to the number of indivi-

dual trees in adjacent plots along a transect, has

proved by this study to be a suitable method for

defining local forest types (conimunities).

Enumcrating cover/abundance ol ail species of a

homogeneous sample plot in rain forest is very

time consuming. It should be noted that the pré¬

sent study was initially not meant lor making a

végétation classification but a general forest

inventory.

The number of 100 m plots (59) may be consi-

dered too small to include the great diversiry of

the forest. More plots are needed and would sur-

ely resuit in clearer différentiation berween the

recognized types and may rcvcal additiona! types.

The size of the 100 m plots (100 X 5 m) might

be questioned. for relevés in rain forest, enume-

rating only trees above a certain dbh larger plots

are often used. h is beyond the scope of this

study to debate the issue of sample plot size and

minimum area. The reader is referred to amongst

others, Hommel (1987, 1990), DE ROUW (1991 )

and DUIVENVOORDF.N & LIES (1995).

The forest typés as distinguished in this survey

show that within the. general fotest classification

of Caballë (1978) for eastern Gabon, vatious

associations can occur. Cabalif! recognized 2

types: a “forêt dense humide sempervirente’’ with

Scyphocephalum ocbocoa , Pycnanthus angolensis,

Pentacletbra eetveldeana , Celtis spp.,

Gilletiodendron piérreanuni, and Gilbertiodendron

dewevrei, a “foret dense à tendance semi-caduci-

foliée" with Pycnanthus angolensis , Pentaelethra

eetveldeana , Terminalia superba and Triphchhon

scleroxylon. These correspond more or less with

Types B and E found in the Minkebé area.

However, in the Minkébé survey Scyphocephalum

ocbocoa, Gilletiodendron pierreanum and

Triplocbiton scleroxylon were not found.

GÉRARD (1960) regarded the Gilbertiodendron

dewevrei forest in the Uele région (Congo) as a

climax végétation, since it was a self regenerating

stable population of this species, EVRARD ( ) 968)

regarded single dominant forest as rhe climax

type since the dominants, besides being able to

regenerate in their own shade, can also invade

mixed moist semi-evergreen rain forest, which is

usually déficient in régénération of its own upper

canopy species. This contrasts with the opinion

of LETOUZEY (1983) who regards the

Gilbertiodendron dewevrei forests as relicts of a

formerly more extensive Gilbertiodendron forest,

tlvat are steadily being invaded by the surroun-

ding mixed forest species, due to natural or

human causes. ÉVRARD and l.FTOUZFY agréé that

possibly recent climatic change has been too

rapid to allow Lhe Gilbertiodendron dewevrei

forest to achieve its maximum potential range

during chose climatic phases of the Pleistocene

most favourable for its expansion. The présent

study reveals that GéRARd’s view on the

Gilbertiondnm forest also applies to the Minkébé

région.

Acknowledgements

Spécial thanks are due to the ficldcrew of Africa

Forest, under the supervision of C. WlLKS, for collec-

ting the data in the field. F. Aleva, L. Oyen, and J.

WtLHlNtlA are gratefiilly atknowltdged for assistance

in processing the data, fl, KLHE5 kindly helped with

sonie of the dtâwing. For assistance in the identifica¬

tion of the herbarium spécimens we thank J J Bos,

F.J. Brltf.i r.R, J.J.FJL de Wii de, C.C.H. Jongkind,

A.J.M. Leeowenberg, R.H.M.J. Lemmens, M.S.M.

SoSEE of the Herbarium Vadense ar Wageningen, and

E. RoBBRECHT at Meise. We are grateful for the

valuable continents on the text given by P.W.F.M.

Hommel and two anonymous referees.

REFERENCES

AL'RREvim E A. 1967-—La forêt primaire des mon¬

tagnes de Bélinga. Biologia Gabonica 3: 95-112.

CabaeLE G. 1978.—Essai sur fa géographie forestière

du Gabon. Adansonia, sér. 2, 17: 425-490.

Duivenvoorden J.F. & Lips J.M. 1995. —A land-

ecological study ofsoils, végétation, and plant diversity

156

ADANSONIA. sér. 3 • 1998 • 20(1)

Floristic inventory in the Minkébé région (NE Gabon)

in Colombian Amdzonia, Tfüpcnbos Sériés 12. The

Tropenbos Foundation, Wâgeningc-n.

ÈVRARD C. 1968.— Recherches écologiques sur le peuple¬

ment forestier ries sois hydromorpbes de la Cuvette cen¬

trale Congolaise. I.N.É.A.C., série scientifique 110,

Congo,

Florence J. & Hiadik A. 1980.—Catalogue des

Phanérogames et des Ptéridophytcs du Nord-Est

du Gabon (Sixième liste). Aaansoniit , sér. 2, 20:

235-253.

GÉRARD Ph. 1960.— Etude écologique de la forêt dense

à Gilbertiodendrnn dewevrei dans la région de l'Uele.

I.N.É.A.C., série scientifique 87, Congo.

H ALLÉ, N. 1964.—Liste des Phanérogames et des

Ptéridophytes des environs de Makokou,

Kemboma et Bélinga. Btolqgia Gahonica 1: 41-46.

Halle N. 1965,—Seconde liste de Phanérogames et

Ptéridophytes du N-E du Gabon (Makokou,

Bélinga et Mékambo). Biologia Gahonica 1: 337-

344.

Hallé N. & Lé Thomas A. 1967.—Troisième liste

de Phanérogames du N-E du Gabon (Makokou,

Bélinga et Mékambo). Biologia Gahonica 3: 113-

120 .

Hali.é N. 6t Ln Thomas A. 1970.—Quatrième liste

de Phanérogames et Ptéridophytes du N-E du

Gabon (Bassin cPIvindo), Biologia Gahonica 6: 131-

138.

Hallé N., Lé Thomas A. & Gazel M. 1967.—Trois

relevés botaniques dans les forets de Bélinga (N.-E.

du Gabon). Biologica Gahonica 3: 3-16.

Hill M.O. 1979.— JWINSfAN: A FORTRAN oro-

gram for diraitging multminate data in an ordered

two-way table by classification of the rndividuais and

attributes. Cornell Univetsity. New York.

Hladik A. & Halle N, 1973.—Catalogue des

Phanérogames du Nord-Est du Gabon (Cinquième

liste). Aaansonia, sér. 2, 13:527-544.

Homme!, P.W.F.M. 1987.— Landscape-eeology of

Ujung-Kulon ( West Java, Indonesia). PhD tfiesis,

pnvately published.

Homméi P.W.F.M. 1990.—A phyrosociological

srudy of a forest in the humid tropics (Ujung

Kulon- West Java, Indonesia. Vegetdtio 89: 39-54,

Kent M. & COKER P. 1992. —Végétation description

and analysis. CRC Press, USA,

LetouZey R. 1983.—Quelques exemples camerou¬

nais de liaison possible entre phénomènes géolo¬

giques et végétation. Botbalia 1-4: 739-744.

RéITSMA JM. 1988.— Végétation forestière du

Gabon/Forest végétation of Gabon. (Technical Sériés

1 ) The Tropenbos Eoundarion, Ede.

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tion in a tropical rain forest. A study of végétation

dyuamics. PhD thesis, privalely published.

Saint Aubin G, de 1963.— La forêt du Gabon.

CTFT, Nogent-sur-Marne.

SCHAiVllNÉH. J.H.J., S l ORI tl DEK A.H.F. & Wl STHOFF

V. 1995 .—De vegetatie vttrt Nederland. Deel 1.

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APPENDIX: NEW RECORDS FOR NORTH EAST GABON

Based on MINKÉBÉ collections présent at Herbarium Vadense (WAG) and fully documented there.

Complété species lise (based on VAN VALKENBURG 1990) published as appendix 2b in STEEL (1992).

Additional collections from december 1990 hâve been included for the présent list.

Acanthaccae

Justicia tenella (Nees) T. Anderson: W 508

Phaulopsis poggei (Lindau) Lindau: W 427

Rhinacantnus virent (Nees) Milne-Redh.: AM 7; W 153

Alismataceae

Ranalisma humile (Kunth) Hutch.: W 144; Wieringa 614

Anacardiaceae

Sorirtdeia sparanoi De Wild.: E 32

Annonaceae

Monanthotaxis klainei Pierre ex Engl. & Diels var. lastoursvillensis (Pellegr.) Verdc.: AM 26

ADANSONIA, sér. 3 • 1998 ■ 20(1)

157

van Valkenburg J.L.C.H., Ketner P. & Wilks C.M.

PachypocLinthium confine Engl. & Diels: C 324, 455; W 65

Uvaria versicolor Pierre ex Engl. & Diels: AM 46

Xylopiaparviflora (A. Rich.) Benth.; C 364, 716

Xylopia pynaertii De Wild.: C 789; E 288

Apocyn.ia.Me

Baisser baillomt Hua: R 141

LandolphiaJbretii Juin.: C 112

Rauvolfîa coffra Sond.: E 26, 347

Rauvolfia letouzeyi Leeuwenb.: E 270

Tabernaemontana eghtndulosa Stapf: W 509

Begoniaceae

Bégonia mildbraedii Gilg: W 200, 436

Bégonia scutifolia Hook. f.: W 221

Bégonia sessilifolia Hook. f.: W 278

Bégonia subicutata De Wild.: AM 77; W 171, 361; Wieringa 539, 581

Bombacaceae

Rhodognaphalon brevicuspe (Sprague) Roberty: A 721

Caesalpiniaceae

Amphimas pterocarpaides Harms; A 72, 467

Anthonalha cl. ferrugfnea (Harms) ). Léonard: C 207

Baphiopsis parviflora Benth. ex Baker: W 57; X 237; Y 6, 13

Cynomclrn sonagaensis Aubrév.: AM 73; E 87; Wieringa 542

Cynometra scblei'bteri Harms: W 496

DanieUia ob/onga Oliv.: A 102

Erytbrophleum suaneolens (Guill, &c Perr.) Brenan: C 686; S 195

Gilbertiodendron ogoouense (Pellegr.) J. Léonard: AM 72; C 33, 68, 89, 118, 283, 367, 371, 387, 484, 492; E

18; VI, 140. 180,231

Guibourtia tessmannii (Harms) J. Léonard: A 77

Julbcrnardia seretii (De Wild.) Troupin: B 83

MonopetaLtnibusemtrdii P. Bamps: A 50; B 43; C 27, 87, 279, 286; E 16, 69, 139, 167, 182; S 41; W438

Monopetakinthm ktesnti Pellegr.: A 360, 609

Oxystigma bachholzii Harms: A 1 ; AM 45; E 2; W 3; Wieringa 540 Z 18

Plagiosiphon anarginatus (Hntcll. & Dalz.) J. Léonard: F. 163, 181 ; S 93; W 591

PUgiosipbtm multijugus (Harms) J. Léonard: C 3,37, 347; E 23, 63, 66. 117; V 2; W 642; X 10

Plagiosiphon multijugus (Harms) J. Léonard var, gracitis Pellegr.: C 32

Stemenocoleus micrantbus IJarms: B 138

Tessrnannia ammutla (Micheli) Harms: B 35, 91; C 8

Tetraberlinia bifolwlata (Harms) Hauman: C 82, 299, 447, 483; E 287

Capparaceae

Cleoma afospinosa IIris: AM 18; W 127

Chryso ba I a naceae

Magnistipulfl zenkeri Engl.: AM 70; E 110, 112, 114

Combretaceae

Combretum concbipetalum Engl. & Diels: E 108, 148

Combreturn demeusci De Wild.: W 409, 502

Combretum bntnalimdes Hutch. & Dalz.: C 99

Combretum multinervium Exell: W 29

Combretum sordtdum Exell: W 227

Strephonema martnit Hook. f.: E 115

Commelinaccae

Commelina afiicana 1„: Wieringa 644

Floscopa africana (P. Beauv.) C.B. Clarke: W 145

Floscnpa glomerata (Willd. ex J.A. & J.H. Schulr.) Hassk.: W 448

Palisota ambigua (P. Beauv.) C.B. Clarke: Wieringa 640

Compositae

Adenostemrna perrortetii DC.: W 115, 333

Ethulia tonysoides L. f.: W 116

Melanthera scandent i,Schum. & Thonn.) Roberty: W 380, 449; Wieringa 584; Y 12

Struchium sparganophora (L.) O. Kuntze: W 344

158

ADANSONIA, sér. 3 • 1998 • 20(1)

Floristic inventory in the Minkébé région (NE Gabon)

Vernonia stellulifera (Benth.) Jeffrey: W 240

Convolvulaceae

Neuropeltis cf. velutina Hallier f.: S 51

Cucurbitaccae

Zehneria gilletii (De Wild.) E. Jeffrey: W 497; Wieringa 659; Y 11,21

Cyperaceae

Bulbostylisfilamentasd (Vahl) C.B. Clarke: W 237

Cyperus compressas L.: W 99

Cyperus digitalus Roxb. subsp. acericomus (Spreng.) Kük.: W 139, 381

Cyperus laxus Lam. subsp. bucbbolzii (Boeck) Lye: W 239

Cyperus remotispieatvs Hooper: W 455

Fuirena umbelUia Roitb,: W 454

Hypolytrum senegalcnse A. Rich.: AM 79; V 104; W 131

Mapania amplivaginata K, Schum.î AM 59; W 432

Dichapetalaceac

Dichapetalum cboristilum Engl.: C 173; R 29; S 5, 87, 219

Dichapetalum lujae De Wild. & T. Durand: S 43, 70

Dichapetalum minutiflorum Engl.: S 61

Dipterocaipaceae

Ancistrocladus barteri Scott-Elliot: R 80, S 231

Ebenaceae

Diospyros alboflavescens { Gürke) F. White: V 95

Diospyrosferrea (Willd.) Bakh.: AM 15; E 142; V 122; W 11, 341

Diospyros sanza-minika A. Chev.: AM 38

Euphorbiaceae

Antidesma venosum Tul.: W 532, 548

Antidesma vogelianum Midi. Arg.: W 120, 150, 188, 505; Wieringa 607; Z 34

Bridelia micrantba (Hochst.) Baill.: C 360

Cleistanthus letouzeyi J. Léonard: E 136, 154; W 377

Erytbrococca anomale (Juss. ex Poir.) Prain: Wieringa 590

Eupborbiagrandifolia (Haw.) Croizat: W 256

Keayodendron briddioides (Mildbr. ex Hutch. & Dalz.) Leandri: A 69; C 227, 261, 449

Maprouncu ufricana Müll. Arg.: C 404

Microdesmispuberula Hook. I. ex Planch.: R 144; S 104

Oldfieldia ufricana Benrh. & Hook. f,: A 660, 663, 712

Pentabraclnon réticulation Müll. Arg.: C 179

Phyllantbus diandrus Pa\: R 138

Pycnocoma corntita Mull. Arg.: W 34

Flacourtiaceae

Flacourtia vogelii Hook. f.: W 480

Homalium abdessamrnadii Aschers. de Schweinf.: W 476

Homalium africanum (Hook. f.) Benth.: AM 56

Oncoba mannii Oliv.: Wieringa 562, 636

Scottellia klaiueana Pierre; A 68

Gramineae

GitaduelLi marantifolia Franch.: R 6

Isachne kiyalensis Robyns: W 456

Leptapsis zeylanica Nées ex Steud.: Wieringa 551

Panicum mueense Vanderyst: Wieringa 573

Guttiferae

Garcinia mannii Oliv.: E 306

Garcinia staudtii Engl.: AM 14; C 566

Psorospemtum lenuijolium Hook. f.: W 262

H ippocrateaceae

Salacia abita De Wild.: R 32; S 150

Salaria leptochtda Fui.: Wieringa 563

Icacinaceac

Alsodeiopsis staudtii Engl.: Wieringa 651

Desmostacbys tenuifolius Oliv. var. tenuifolius-, Wieringa 645

ADANSONIA, sér. 3 • 1998 • 20(1)

159

van Valkenburg J.L.C.H., Ketner P. & Wilks C.M.

Icacina tmtnnii Oliv.: R 146; S 73

Pyrenacantha vogeliana Baill.: S 145

Irvingiaceae

Desbordesta glaucescens (Engl.) Tiegh.: A 402

Irvingia exct-lsa Mildbr.: A 472

Klainedoxa microphylla (Pellegr.) A. Gentry: W 299a

Klainedoxa trillesii Pierre ex Tiegh.: A 543

Labiatae

Hoslundia opposita Vahl: W 568

Platostoma ajricanum P. Beauv.: W 358

Solenostemon matinii (Hook. f.) Baker: W 447

Liliaceae

Chlorophytum laxum R. Br.: W 149

Chlorophytum orchidastrum Lindl.: W 387

Dracnena mannii Baker: A 289

Dracaena viridiflora Engl, ex Krausse: D 63; W 255

Linaceae

Hugonia afzelii R. Br. ex Planch.: W 283

Loganiaceae

Strychnos bonnet De Wild.: V 55

Sttycbrws congobma Gilg: R 166

Strychnos staudtii Giig: V' 183

Strychnos tchibangensis Pellegr.: W 396

Loranthaceae

Globimetula opaca (Sprague) Danser: W 596, 605

Phragmantheru capitata (Sprcng.) Balle: Wieringa 593

Tapinarlthus platyphyllus (Hochsc. ex A. Rich.) Danser: W 544

Viscum congvleme De Wild.: AM 64; F 4; W 68

Luxemburgiaceae

Testuleagabonensis Pellegr.: A 801; C 330

Malvaccae

Hibiscus rostellatsu Gnill. & Perr.: W 126

Marantaeeae

Halopegia azurea (K. Schum.) K. Scbum.: W 151; Wieringa 545

Marantochloa etmgensis (K. Schum.) J. Léonard & Mull.: V 188

Marantochlou leucantha (K. Schum.) var. leucantha: Wieringa 585

Marantochloapurpurea (Ridl.) Milne-Rcdh.: AM 1; W 313; X 11

Sarcophryniutn brachystachys (Benth.) K. Schum.: W 17

Melastomaraceae

Calvoa orientalis Taubert: W 243

Guyonia ciliata Hook. f.: W 419

Memecylon vmde I lurch. & Dalz.: W 343

Ochthocharis dicellandroid.es (Gilg) C. Hansen & Wickens: W 452

Tristemma littorale Benth. subsp. biafraniim}ac<\.-Yi\.\ AM 43; W 158

Meliaceae

Carapa proccra DC.: C 77

Entandrophragma congoense (De Wild.) A. Chev.: B 243, 313

Entandrophragma cylmdricum (Sprague) Sprague: A 63; B 40; C 439

Guarea cedrata (A. Chev.) Pellegr.: B 69; C 256

Guarea thomsonii Sprague & Hutch.: A 469; B 74

Trichitia monadelpha (Thonn.) J.J. de Wilde: E 104, 361; W 660; X 236; Z 25

Trichilia tessnuinvii Harms: A 21, 634; X 236; Z 25

Trichilia wclwitschii C. DC.: AM 31; C 687; E 268

Turraeantbus africanta (Welw. ex C. DC.) Pellegr,: A 435

Mimosaceae

Albizia zygia (DC.) J.F. Macbr.: W 75

Newtonia dupdrquetiana (Baill.) Keay: W 526, 550

Parkia filicoidea Welw. ex Oliv.: A 117, 397

160

ADANSONIA, sér. 3 • 1998 • 20(1)

Floristic inventory in the Minkébé région (NE Gabon)

Moraceae

Antiaris africana Engl.: A 643

Dorstenia mannii Hook. f.: W 223, 274, 637, 645

Ficus adolfi-fiiderici Mildbr.: V 62

Ficuspseudomangifera Hutch.: W 8

Milicia excelsa (Welw.) C.C. Berg: A 78; C 700

Nymphaeaceae

Nytnphaea lotus L.: W 606

Ochnaceae

Ouratea ({[finis (Hook. f.) Engl.: W 122, 352; X 72

Ouratea calarttha Gilg: W 95, 469; X 146

Ouratea congesta (Oliv.) Engl.: AM 66; R 99; S 171 ; W 41, 85, 94

Ouratea elongata (Oliv.) Engl.: W 204; Wieringa 587

Olacaceae

Strombosia schefflcri Engl.: A 104; C 262, 335; E 201 ; V 93; W 491

Orchidaceae

Bulbopbyllum cocointtm Lindl.t Wieringa 580

Genyorchis apetala (Lindl.) J.J. Vermeulen: Wieringa 538

Papilionaceae

Airyantba scbioeinjurthii (Taub.) Brummitt: S 132

Baphia buettneri Harms subsp. hylophila (Harms) Soladoye: AM 9; W 88

Baphia leptostemtna Baill.: C 76

Baphiapubescetis Hook. f.: AM 35; C 211,423; F 6; W 58

Baphiastrum braclrycarpum Harms: S 44

Bapbiopsts parviûora Benth. ex Baker: Wieringa 547

Millettia bipindensis Harms: AM 49, 69; C 352; E 217;W 152, 493; X 100; Z 30

Pontederiaceae

Heterantbera callifolia Rchb. ex Kunth: AM 24; W 141

Pteridophyrac

Antrophyurn mannianum Hook.: V 185; W 195

Asplénium jaunclense Hieron.: V 136; W 347

Blotiella currori (Hook.) Tryon: W 471

Diplazium sammalii (Kuhn) C. Chr.: W 146; Wieringa 619

Polypodium owariense Desv.: AM 12; W 348; X 108

Tricbomanes balletrdianum Alston: V 72

Trichonuwes mannii Hook.: V 73; W 369

Triplophyllum gaboneme Holttum; R 4; S 19

Triplophyllum vogelii (Hook.) Holttum: W 228, 446

Rhamnaccac

Maesopsis eminii Engl.: E 354

Rubiaceae

Belatwphora arborescent Hoyle: W 285

Commitheca liebrechtsiana (De Wild. & T. Durand) Bremek.: W 305, 478

Didimosalpinx abbeokutae (Hiern) Kcay: E 179

Geophita involucrata Scbweinf. ex Hiern: S 31; W 417

Hymenoeoleus globulifer Robbrechr: W 166

Hymenocoleus hirsutus (Benth.) Robbrecht: R 36; S 72, 109; W 279, 4l8bis, 463

Hymenocoleus nervopilosus Robbrecht var. orientait! Robbrecht: W 315, 418

Hymenocoleussubipccactumha (K. Schum.) Robbrecht: AM 61; R 5; W 202

Ixoragutneensis Benth.: W71, 197;X 186

Mitragyna stipulosa (DC.) O. Kuntze: A 604

Oldemandia goreemis (DC.) Summerh.: W 444

Pseudosabicea medusula (Wernham) N. Halle: AM 29; W 55, 163

Rothmannia whitfieldii (Lindl.) Dandy: E 338

Sabicea dinklagei K. Schum.: W 378

Tricalysia vaclensis Robbrecht: R 128, 232; W 420

Sapindaceac

Allophylus cobbe (L.) Raeusch.: R 254; W 483

Deinbollia maxima Gilg: W 400

ADANSONIA, sér. 3 • 1998 • 20(1)

161

van Valkenburg J.L.C.H., Ketner P. & Wilks C.M.

Sapotaceae

Gambeya africana (Don ex Baker) Pierre: W 295

Manilkara argentea Pierre ex Dubard: W 119

TieghemelLi africana Pierre: A 659

Tridesmostemon omphalocarpoid.es Engl.: A 640

Scytopetalaceae

Oubanguia bmrifolia (Pierre) Tiegh.: Y 19

Simaroubaceae

Odyenda gabonensis (Pierre) Engl.: C 318, 782; S 139

Sterculiaceae

Eribroma oblonga (Mast.) Bodard: A 25; C 194

Leptonychia echinocarpa K. Schum.: Wieringa 621

Leptonychia lasiogytie K. Schum.: F 5

Leptonychia multiflora K. Schum.: AM 37; R 64; W 13, 13bis, 49, 110

Sterculia subviolacea K. Schum.: W 1

Thymelacaceac

Dicranolepis baertsiana De Wild. &: T. Durand: W 63, 181

Dicranolepis vestita Engl.: AM 8, 23; W 62, 101

Tiliaceae

Duboscia viridiflora (K. Schum.) Mildbr.: C 186

Grewiapinnatifida Mast.: C 54; E 165

Ulmaceae

Celtis adolfi-friderici Engl.: A 126; AM 55

Urticaceae

Laportea ovalifolia (Schum. & Thonn.) Chev.: W 51,401

Violaceae

Rinorea cerasifolia M. Brandt: W 102, 160, 277, 339

Rinorea karnerunensis Engl.: D 16

Rinorea longicuspis Engi.: Wieringa 546

Rinorea müdbraedii M. Brandt: Wieringa 561

Rinorea subsessilis M. Brandt: AM 2; C 618; D 6; R 8; W 467

Rinorea welwitschii (Oliv.) O. Kuntze: C 83; R 239; W 258

Vochysiaceae

Erismadelphus exsul Mildbr.: A 184

Zingiberaceae

Aframomum limbatum (Oliv. & Hanb.) K. Schum.: W 443; Wieringa 554

Renealmia cincinnata (K. Schum.) Baker: W 92

Explanation of préfixés in Minkébé collection: A, B: voucher specimen transect trees > 70 cm dbh; C, E: voucher specimen tran-

sect trees > 10 cm dbh; R, S, V, X, Y, Z: voucher specimens subplots; AM: André Moungazi; D: Dibata; F: Mondjo, Obianq; W:

Wilks.

Manuscript received 21 June 1997;

revised version accepted 26 February 1998.

162

ADANSONIA, sér. 3 • 1998 • 20(1)

Sylvichadsia, a new genus

of Leguminosae-Papilionoideae-Millettieae

endemic to Madagascar

Jean-Noël LABAT

Laboratoire de Phanérogamie, Muséum national d'Histoire naturelle,

16 rue Buffon, 75005 Paris, France.

Iabat@mnhn.fr

David J. DU PUY

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, England.

d.dupuy@rbgkew.org.uk

KEYWORDS

Leguminosae,

PapMionoideae,

Sylvichadsia ,

Madagascar.

ABSTRACT

Sylvichadsia , a new genus in the tribe Millettieae (Leguminosae-

Papilionoideae) is described. It includes four species, ail endemic to

Madagascar (5. macrophylla (R. Vig.) Du Puy & Labat, S. grandifolia (R.

Vig.) Du Puy 6c Labat, S. grandidieri (Baill.) Du Puy & Labat, and S. perrie-

ri (R. Vig.) Du Puy & Labat).

MOTS CLÉS

Leguminosae,

Papilionoideae,

Sylvichadsia,

Madagascar.

RÉSUMÉ

Sylvichadsia, un nouveau genre, de la tribu des Millettieae (Leguminosae-

Papilionoideae) est décrit. Endémique de Madagascar, il renferme quatre

espèces (S. macrophylla (R. Vig.) Du Puy & Labat, S. grandifolia (R. Vig.)

Du Puy & Labat, S. grandidieri (Baill.) Du Puy & Labat, et S. perrieri (R.

Vig.) Du Puy & Labat).

ADANSONIA, sér. 3 • 1998 • 20(1)

163

Labat J.-N. & Du Puy D.J.

The tribe Millettieae is one of the largest tribes

of Papilionoid legumeS, aJong with the

Phaseoleae and the Sophoreae (POLHILL 1981); it

is diverse and contains numeruus species in

Madagascar. In order to produce a complété

treatmeitt of the Leguminosae-Papilionoideae for

Madagascar and the Comoros, we hâve srudied

ail of the collections of Millertieae from rhis

région held in G. K, MO. P. TAN. THF and

WAG. This research has shown that Madagascar

is an important and old centre of diversity for

this tribe (LABAT 1996), with 77 spécifie or sub-

specific taxa (68 of them endemic) in 10 généra

(5 endemic). Pyranthus has already been revised

for Madagascar (Du PUY & Labat 1995) and

new spccies and new combinations in MiUettia

Wight & Arnott and Pongamiapsis R. Vig.

(Labat & Du Puy 1995) hâve been published.

GEESINK (1981 ) isolated a large group of généra

within the Millettieae (as “Tephrqsieae”) with

assumed inter-relationships, centred around the

large genus Tephrosia Pers. (with over 400 spe¬

cies). lt included, in addition to Tephrosia, the

généra Mundulea (DC.) Benth. and Chadsia

Bojer, both of them confined to Madagascar

with the exception of Mundulea sericea fWilld.)

A. Chev. (which is a widespread subspeaes occurr-

ingfrom Africa to Southern Asia). GlTMNK (1984)

considered rhat rhe position of Mundulea needed

révision, especially with regard to Tephrosia. It

appears that Mundulea is heterogeneous, and we

hâve described a new genus nanled Pyranthus Du

Puy & Labat, which includes 5 species originally

described in Mundulea or Tephrosia , and a single

new species (Du PUY & LaüAT 1995).

Similarly, Mundulea microphylla R. Vig. must

be excluded from the genus Mundulea as well as

two species described in the genus Chadsia ,

C. grandifolia and C. gmndidieri, since ail three

species exhibit characters differing from both of

these généra: the présence of bracteoles, a distinct

hypanthium, a truncate calyx and a cauliflorous

habit. In his révision of the genus Strongylodon,

Huang (1991) excluded from this genus a very

little known species described bÿ VlGUiER (1952)

as Strongylodon perrieri, on the basis of the leaves

lacking stipels, the absence of a discoid nectary

and other floral and végétative characters,

without making any suggestion as to its correct

classification. It appears that a new genus ende¬

mic to Madagascar can be recognised in this

complex to accommodate these species: it is des¬

cribed here as Sylvichadsia.

Sylvichadsia is characterised by its cauliflorous

habit, rhe presence of bracteoles on the pedicels,

its truncate calyx, its wide and distinct hypan¬

thium, its stipitate ovary and its pods which are

déhiscent, splitting into 2 spiralling valves. The

presence ol a hypanthium and bracteoles are

considered to be primitive features in the

Millettieae, and studies of the évolution of this

tiibe should include this genus. It has already

been demonstrated that the généra Phylloxyllon

and Vaughania, also endemic to Madagascar, exi-

bit some primitive characteristics within the tribe

Indigofereac (Du Puy et al. 1994, 1995), and, in

cladistk analyses of this tribe, these généra form

basal branches of the cladogram (SCHRIRE 1995).

Sylvichadsia may be similarly basal within the

Millettieae. The torm of the flowers (especially

the strongly beaked keel and evenly curved

androecium and gynoecium) and the pods of

Sylvichadsia grandi folia and V. gmndidieri closely

rcsemble those of Chadsia , but they differ in the

suberect standard pctal and the other characters

listed above. The resemblance between the flo¬

wers of these two species and those of Chadsia

can be interpreted as a convergence or parallel

évolution of the flowers linked with adaptations

to a bird pollinarion syndrome.

The flowers of S. macropljylla are white in colour

and niost closely resemble those of some Millettia

species, with a straight androecium and gynoe¬

cium sharply upturned at the tip, and a blunt keel

tip. Howcver, this species shares ail of the generic

characteristics listed above (although the pods are

not known); its flowers appear diffèrent from

those of the three other species, but ail the différ¬

ences are linked with the biologtcal spécialisation

of those other species to bird pollinarion, and S.

rnacrophylla cannot be excluded from Sylvichadsia.

Its hairy androecium is extremely unusual.

Sylvichadsia perrieri is a liane with trifoliolate

164

ADANSONIA, sér. 3 • 1998 • 20(1)

Sylvichadsia (Leguminosae)

leaves, the Flowers are produced in long pseudo-

racemes and the keel rip has a rather blunt apex.

Its habit and leaves are very difFerent from the

other species, but the flowers resemble those of

S. grandifolia and S. grandidieri. and they also

retain the generic characteristics listed above.

Although this species was originally described in

the tribe Phaseoleae, it is included here in the

genus Sylvichadsia. The trifoliolate leaves are ver)'

unusual in the Millertieae, although they can be

considered as a réduction From imparipinnate

leaves. This character occurs in many higher

tribes ol the Papilionoideae, but it also appears in

the tribe Miilettieae, in Disynstemon R. Vig., a

monotypic genus endetnic to Madagascar. The

vine habit can also be considered as a parallelism

appearing in many généra oF the tribes

Sophoreae and Miilettieae, such as in the généra

Bapbia or Millettia.

The generic name is derived From “silva/sylva”

meaning “fbrest" and Chu dsi a. another genus in

the Miilettieae endemic to Madagascar, which

has similarly beaked and orange-coloured Flowers

(especially resembling those ol S. grandifolia and

5. grandidieri). Sylvichadsia occurs in the humid,

eastern rainforests, while Chadsia occurs in the

seasonally dry, deciduous végétation oF western

and southem Madagascar.

SYLVICHADSIA Du Puy & Labat, gen. nov.

Arbor, jrutex vd liana lignard, retaper niuliftora. folia

imparipinnata vd trifolioUna ; stipdlac 0. ltifhresctntiae

in trunco vd in ramis crassisümis dhpôskae, fasciculatae,

racemosae. pseudoraeemosae vel patii.ad.atae, bmcteolae

persistentes in media vel in ima parte pedicelli dispositae.

Flores rnagnae vd médiocres. 17-45 mm longae, viola-

ceae, rttbrae vel albae. Calyx atpularis tel carnpauulatus,

apice truncato edentato vel 4-5 prominentih minutis

proviso; hypanthium latum eonspicitumque. Vcxdlum

erectum vel suberecutm. Antberac in longitudinem

aequantes. Ovanum glabrum, breviter vtllosum vel

pubescens, brève stipitatum, ovulis multis; srigmantm

minutum obliquum.

TYPUS.— Sylvichadsia grandifolia (R. Vig.) Du Puy

& Labat (= Chadsia grandifolia R. Vig.).

Cauliflorous trees, shrubs or woody lianes (in S.

perrieri); deciduous or evergreen. Leaves large,

usually dustered on the twig or stem tips, iinpa-

ripinnate wirh paired leaflets or pinnately trifo-

liolate (in S. perrieri ), glabrous or almost so,

glabrescent; stipels absent. Inflorescences from

w.irty growths on the trunk and main branches,

the flowers in whorls, racemes, pseudoracemes or

panicles; pedicels with a pair of small, persistent

bracteoles usually near the middle or in the lower

halF. Flowers large or medium-sized, 17-45 mm

long, purplish to reddish or white (in S. rnacro-

phylla). the standard with an irregular, whirish

basal patch. Calyx cup-shaped to bell-shaped,

sometinies narrowly so, not hooded, the apex

truncate and without teeth or the teeth reduced

to 5 minute points; hypanthium wide and dis¬

tinct. Standard petal subcrect to ercct, glabrous

or subglabrous behind; wing petals shorter than

or as long as and appressed to the keel; keel

either extended at the tip into a distinct, narrow

bealc, or blunt and not beaked (in S. manvphyl-

ld)\ androeciutn and gynoecium shallowly cur\'ed

to the tip or strongly upeurved apically. Stamens

fused into a curved or straight sheath, rhe outside

rarely shortly hairy (in 5. inatrophylla ); anthers

equal. Ovary shortly sripitate, rnulti-ovular; style

terete or flattened, short-hairy along the inner

surface or glabrous (in S. perrieri ); stigma minu¬

te. Pods stipitate, long and narrowly oblong, bea¬

ked, splitting into 2 spiralling valves, with several

distantly spaced seeds. SeecLs not known.

A genus oF 4 rare to very rare species confined

to the more humid Forests of eastern and nor-

thern Madagascar. Sylvichadsia can be divided

into 3 natural groups;

Group 1: Sjnall trees or shrubs; leaves multifo-

holate; flowers in fascicles, short racemes, or few-

branched panicles; flowers large, 30-45 mm long,

red or purplish; keel tip extended into an acute

beak; staminal s'heath glabrous; androecium and

gynoecium evenly curved along their length.—1,

S. grandifolia ; 2. S. grandidieri.

Group 2: Shrub; leaves multifoliolate; pseudo¬

racemes short and dense, with numerous flowers

in closely spaced clusters along the inflorescence

axis; flowers small, 17-20 mm long, white; keel

tip rounded, not extended into a beak; staminal

ADANSONIA, sér. 3 ■ 1998 • 20(1)

165

Labat J.-N. & Du Puy D.J.

sheath hairy; androecium and gynoecium

straight and sharply upturned at the tip.—3, S.

macrophylla.

Group 3: Liane; leaves 3-foliolate; flowers in

long and lax pseudoracemes; flowers medium

si/ed, 22-24 mm long, brigKt scarlet red; keel tip

obtuse, not extended into a long beak; staminal

sheath glabrous; androecium and gynoecium

evenly curved along their lcngrh.—4, S. perrieri.

Key to Sylvichadsia

1. Shrub or tree; leaves with 5-23 leaflets; flowers in fascicles or in short racemes, panicles or dense pseudora¬

cemes up to U) cm lone; keel tip extended into an acute beak, or if rnore roundcd then flower white

(otherwise red or purplish) .......2

1’. Liane; leaves with 3 leaflets only; flowers in pseudoracemes ca. 45 cm long; keel tip obtuse, not extended

into a beak (flowers bright red) ....4. S. perrieri

2. Flowers smaller, 17-20 mm long, white; keel tip rounded, not extended into a beak; staminal sheath hairy;

pseudoracemes dense, with numerous flowers in closely spaced clusters along the inflorescence axis ...

......... 3. S. macrophylla

2’. Flowers larger, 30-45 mnt long, red or purplish; keel tip extended into an acute beak; staminal sheath gla¬

brous; inflorescences not as above, the flowers in fascicles, short racemes or few-branched panicles .3

3. Calyx cup-shaped, as wide as long or wider; standard 17-30 mm wide; pedicels short, 5-10 mm long;

leaves with 5-9 leallets .......... 1. S. grandifolia

3’. Calyx tubular, almost rwice as long as wide; standard 14-16 mm wide; pedicels 10-20 mm long; leaves

with 9-23 leaflets.2. S. grandidicri

1. Sylvichadsia grandifolia (R. Vig.) Du Puy &

Labat, comb. nov.

Chadsia grandifolia R. Vig., Not. Syst., Paris 14: 70

(1950).—Type: Perrier de la Bâthie 4110,

Sambirano: forêt du Sambirano, Oct. 1908, fl.

(lecto-, P; isolccro-, P: chosen here).

Mundulea macrophylla R. Vig., Not. Syst., Paris 14:

67 (1950), pro parte lectotypc excl.

A small, evergreen tree or large shrub 4-12 m

tall, cauliflorous; trunk reaching 30 cm in dia-

meter, with smooth, greyish bark, releasing a

strong, acrid odour when damaged. Leaves in

terminal clusters, large, 15-45 cm long, with 5-9

leaflets (2-4 pairs)! leaflets large, elliptic to

oblong-elliptic, 6-19 X 2.5-9 cm, the apex acu-

rninate and finally obtuse, glabrous, subcoria-

ceous. Flowers in short racemes or few-branched

panicles Ü.5-6(-10) cm long with up to ca. 15(-

20) flowers, produced from warty outgrowths

mostly ou the trunk or older branches; pedicels

short, 5-10 mm long, with a pair ot small brac-

teoles near the middle or towards the calyx.

Flowers 33-45 mm long, deep red or purplish

with an irregular, whitish patch at the base of the

standard. Calyx wide, cup-shaped, truncate, as

wide as long or wider, 7-10 mm long, pubescent

ro glabrous except for the rim; teeth absent or

reduced to 5 very small points on the calyx rim.

Standard liinb suberect or pointing backwards,

widely ovate, 30-40 X 17-30 mm, the apex acute,

glabrous, the claw fleshy; wings 6-8 mm wide,

slightly shorter than the keel; keel 8-1 1 mm

wide, 45-60 mm long (measured from the base

of the claw, around the lower tnargin to the tip

of the beak), sickle-shaped, shallowly but evenly

curved along its length, with a short, blunt,

upeurved beak. Staminal sheath 25-35 mm long,

evenly curved along its length, glabrous. Ovary

shortly stipitare; style- thinly pubescent along the

inner surface. Fruit up to 20 cm long, 15-22 mm

wide, fiat, apprcsscd-pubescent with short, rust-

coloured hairs.—Fig. 1A-G.

V1GUIE.R cited Perrier de la Bâthie 18176 and

18190 as syntypes of Sylvichadsia macrophylla (as

Mimdulea macrophylla). Thcse two specimens

were collected at almost the same locality.

Howcver, the inflorescences and flower shapes

are very different, and Perrier de la Bâthie 18190

166

ADANSONIA, sér. 3 • 1998 • 20(1)

Sylvichadsia (Leguminosae)

Fig. 1.—A-G, Sylvichadsia grandifolla: A, inflorescence (produced from the trunk); B, calyx (longitudinal section showing the

hypanthium); C, calyx exterior; D, androecium and style; E, gynoecium; F, mature leaf; G, pod.—H-L, Sylvichadsia macrophylla: H,

inflorescence (produced from the trunk); I, calyx (longitudinal section showing the hypanthium); J, calyx exterior; K, androecium and

style (showing the pubescent staminal sheath); L, gynoecium. (A-G, Service Forestier 9237-SF-, H-L, Perrier de la Bâthie 18176 ).—

Drawn by Angela Beaumont.

ADANSONIA, sér. 3 • 1998 • 20(1)

167

Labat J.-N. & Du Puy D.J.

most closely resembles Sylvichadsia grandifolia

(inflorescence a short spur, flowers large with a

beaked keel and wings shorter than thekeel, style

glabrous, staminai sheath 22-23 mm long and

evenly eurved along its length). Nevertheless, the

whitish flower colour, the presence of a very few

hairs on the staminal sheath and the more sou-

therly locality in the samc area as Sylvichadsia

macrophylld , are anomalous and indicate that fur-

ther collections from eastern Madagascar are nee-

ded to clarifÿ the species délimitations and

distributions in this genus.

Distribution. —NE & N Madagascar, near

Antalaha, Andapa, Sambava and Antsirabe [du

nord] (300-600 nt), and on Montagne d’Ambre

(600-1000 m).

HABITAT. —ln shaded areas in humid evergreen

forest, often near screams or rivers, on acidic

(basaltic) soils, at 300-1000 m altitude.

FLOWER1NG TIME.—Scptember-November.

LOCAL uses. —The wood is soft, brittle, white,

with an acrid odour when fresh; ail parts of the

plant release potent fîsh toxins upon décomposi¬

tion.

VERNACUI AR NAMF.—Fanamo (NE).

Material examiner. — Madagascar: Homolle 198,

s.loc., s.d., fl. (P); Humbert & Cap won 24218, valle'e

de l’Andalangy, affluent de l’Androranga (bassin de la

Bemarivo du Nord-Est), 300-500 m, 12/14 Nov.

1950, j.fr. (G, K, MO, P, WAC); Perner de la Bâthie

18190, Est: bassin inf, du Mangoro, Oct. 1927, fl.,

j.fr. (K, P); Perrier de la Bâthie 18860 , Centre (Nord):

Montagne d’Ambre. abondant surtout à la base Est du

Pic Baaens, C3. 1000 m, Nov. 1932, fl., fr. (K, MO,

P); Réserves Naturelles 8299 RN, Doany, Andapa, 11

Oct. 1956, fl. (K, P, TEF) ; Réserves Naturelles 9087

RN Sajy, Anrongondriha, Canton Ambohimttsinjo,

Dt. Sambava, 18 Sep. 1957. fl. (P, TEF); Service

Forestier 203 R 152 , Roussette, JB 19, Joffreville,

Diégo-Suarcz, 630 tn, 6 Apr 1955, st. (K, P): Service

Forestier 811 SF, bord d’affluent de l’Andalangy

(affluent de l’Androranga), Di. de Sambava, 350-

500 m, 12 Nov. 1950, fr. (G, K, MO, P, PRE, TEF);

Service Forestier 9237 SF Capuron. Est: Forêt

d’Andrakaraka (au S-SW d’Antalaha), 20/24 Sep.

1954, fl. (K, P, TEF); Service Forestier 11283 SF

Capuron, Centre (Nord): massif de la Montagne

d’Ambre, 800-900 m. 8/14 Oct. J9M, H. (B. BR, G,

K, MO, P, TEF, WAG); Service Forestier 24906 SF

Capuron , Est (Nord): Analamanara, près du village de

Tsaratanana, entre Antsirabe-Nord et Sambava, 24

Oct. 1966, fr, (K, P, TEF); Service Forestier 24962 SF

Capuron , Est (Nord): table basaltique

d Ambanitazana, près d’Andrapengy, au N

d’Antalaha, 22 Oct. 1966, fl. (K, P, TF.F); Service

Forestier 29191 SF Capuron, Centre (Nord) jusqu’aux

confins de l’Ouest (Nord): massif de la Montagne

d’Ambre, crête entre les bassins de la rivière des Makis

et de la rivière d’Amkazobe, 600-800 m, 26/27 May

1970, Ir. (P, TEF); Ursch 204, Forêt d'Ambre, Diégo-

Suarez, s.d., fl. (K, P).

2. Sylvichadsia grandidieri (Baill.) Du Puy &

Labat, comb. nov.

Chadsia grandidieri Baill., Bull. Mens. Soc. Linn. Paris

1: 391 (1883), pro parte lectotype incl, ( Grandidier

5, also cited by Baii i on, is a specimen of Chadsia

flammea), —Type: Lantz 22, Ambakoubé

[Ambakobe], 960-1000 m, 16 june (reçu le 22 Feb.

1882), fl. (lecto-, P, chosen here).

A small, evergreen tree or shruh 2.5-3 m tall,

cauliflorous. Leaves in terminal clusters, large,

35-52 cm long, with 9-23 leaflets (4-11 pairs);

ieailecs narrowly elliptic, 6-16 X 1.5-4 cm, the

apex acuminate and final!y obtuse, completely

glabrous. Flowers in fascicles or on very short

spurs up to 5 mm long from warty outgrowths

on the trunk or older branches; pedicels 10-

20 mm long with a pair of minute bracteoles

near the middle. Flowers 30-35 mm long, red.

Calyx tubular, truncate, aimost twice as long as

wide, 7-9 mm long, very sparsely appressed-

pubescent, tecrh aimost absent. Standard limb

suberect or pointing backwards (folded ca. 5 mm

in front of the calyx tip), ovate, ca 30 x 14-

16 mm, the apex acute, glabrous except for a few

short hairs near rhe rip; wings 4-6 mm wide,

slightly shorter than rhe keel; kee! narrow, 5-

8 mm wide, 42-48 mm long (measured from the

base of the claw and around the lower margin to

the tip of the keel), sicklc-shapcd, shallowly and

evenly eurved, with a short, acute, upturned

beak. Staminal sheath ca. 30 mm long, evenly

eurved, glabrous. Ovary shortly stipitate; style

thinly pubescent along the inner surface. Fruit

not known.—Fig. 2H-M,

DISTRIBUTION. —SE Madagascar, onty known

from 2 localities (20-21 km S of Farafangana on

the route to Réserve Spéciale de Manombo [0-

168

ADANSONIA. sêr. 3 • 1998 • 20(1)

Sylvichadsia (Leguminosae)

Fig. 2. — A-G. Sylvichadsia perrieri A, mature leaf and inflorescence; B. floral bud; C, mature flower; D, calyx (longitudinal section

showing the hypanthium); E, calyx exterior; F, androecium; G, gynoecium.—H-M, Sylvichadsia grandidieri H, mature leaf; I. inflo¬

rescence (produced from older branches); J, mature flower; K, calyx (longitudinal section showing the hypanthium), L, calyx exterior;

M, androecium; N, gynoecium. (A-G, Perrier de la Bâthie 18194-, H-J, Lantz 22\ K-N, Service Forestier 9207-SF). —Drawn by

Dominique Storez.

ADANSONIA, sér. 3 ■ 1998 • 20(1)

169

Labat J.-N. & Du Puy D.J.

130 m altitude], and “Ambakobe” [perhaps

Ambatobe near Farafangana?] at 900-1000 m).

HABI TAT.—In humid, evergreen forest on Iate-

ritic (basaltic) soil or sand, at sea level to 1000 m

altitude.

FloWERING TIME. —Only recorded in June.

MaTERIAL EXAMINEE).—MADAGASCAR: Service

Forestier 9207 SF Captiron, Forêt de Manombo, au S

de Farafangana, 26 June 1954, fl. (K, P); Service

Forestier 23620 SF Capuron , Est: au sud de

Farafangana, route de Manombo, aux P.K. 20-21,

14/17 Oct. 1964, fl. (P, TEF).

3. Sylvichadsia macrophylla (R. Vig.) Du Puy

& Labat, cotnb. nov.

Munduleu macrophylla R. Vig., Not, Syst„ Paris 14:

67 (1950), pro parte lectotype incl. (sec also

Sylvichadsia qrandifolid ).— t ype: Perrier de la Bâthie

18176, Est: Bassin inf. du Mangoro, ea. 200 m, Oct.

1927, fl. (lecto-, P; isolecto-, K, P; chosen here).

A shrub 3-4 m tall, cauliflorous. Leaves in ter¬

minal clusters, large, 1 5-25 cm long, with (5-)7-

9 leaflets (usually 3 or 4 pairs); leaflets large,

elliptic, 5-11 X 2-4.5 cm, tlie apex shortly acu-

minate and finally obtuse, thinly appressed-hairy

above and beneath at first, glabrescent, becoming

coriaceous. Inflorescences dense pseudoracemes

up to ca. 10 cm or more long (perhaps longer as

the inflorescence matures) produced from the

main woody branches, the flowers numerous in

dense, closely spaced clusters on short spurs

along the main axis; pedicels ca. 8 mm long,

with 2 small bracteoles near the middle, Flowers

ca. 17-20 mm long, white. Calyx wide, cup-sha-

ped, truncate, as wide as long or wider when

mature, ca. 6 mm long, thinly pubescent; teeth

reduced to 5 very small points. Standard limb

erect, widely elliptic, ca. 15-18 X 12-17 mm, gla-

brous; wings ca. 4.5 mm wide, as long as the

keel; keel ca. 5.5 mm wide, ca. 20 mm long,

slightly curved, the apex rounded (not beaked).

Staminal sheath straight, ca. 14-16 mm long, the

exterior covered in short, appressed hairs, the

apex and the free portion of the filaments upeur-

ved. Ovary shortly stipitate; style short-hairy.

Fruit not known.—Fig. 1H-L.

The flowers of this species resemble those of

Millettia or Mundulea (keel short, blunt, not

extended into a sharp beak, the ovary and stami¬

nal tube straight, with the style and free portion

of the filaments strongly upeurved).

Neverrheless, the cauliflorous habit, the leaf and

leaflet size, the presence of bracteoles and an

hypanthium, the truncate calyx and the stipitate

ovary closely resemble Sylvichadsia grandifolia,

and there can be no doubt that they are closely

related.

DISTRIBUTION. —E Madagascar, known only

from the type locality in the lower Mangoro

River basin (ca. 20°S-48°30’E).

HABITAT. —Remnants of humid, evergreen, eas-

tern forest, at ca. 200 m altitude.

FloweriNG TIME.— October.

4, Sylvichadsia perrieri (R. Vig.) Du Puy &

l.abat, comb. nov.

Strongylodon peirieri R. Vig., Not. Syst., Paris 14: 175

(1952).— Type: Perrier de la Bâthie 18194, Est:

Bassin inf du Mangoro, ca. 200 m, Ocr. 1927, fl.

(holn-, P; iso-, P).

A woody liane; stems short-hairy when young,

soon glabrescent; stipels absent. Leaves pinnately

3-folioIate; leaflets (25-145-90 X (l5-)25-55 mm,

rounded at the base, shortly tapering apically,

glabrous and glossy above, with a few hairs espe-

cially on the veins beneath, becoming entirely

glabrous, coriaceous; stipules narrowly triangular,

becoming rigid. Pseudoracemes ca. 45 cm long,

the axes finely hairy with short, erect, rusty or

red-brown hairs, the flowers numerous mosdy in

clusters of 3 on swollcn nodes along the axis;

pedicels 10-15 mm long, hairy with a pair of

minute bracteoles inserted in the lower half.

Flowers 22-24 mm long, bright scarlet-red inclu-

ding the calyx. Calyx bell-shaped, 5-6 mm long,

thinly pubescent outside, pubescent wirhin, with

a distinct hypanthium; teeth 4, very small, trian-

gular, the upper tooth notched. Standard limb

erect, oblong-elliptic ca. 19-20 X 12-13 mm,

notched apically, glabrous, auriculate but

without basal appendages, with a claw' ca. 6 mm

long; wings as long as the keel, the limb narrowly

170

ADANSONIA, sér. 3 • 1998 • 20(1)

Sylvichadsia (Leguminosae)

oblong, ca. 18 X 4.5 mm, auriculate basally, with

pubescent margins towards the base; keel curved,

ca. 19 X 6 mm, obtuse (not beaked), auriculate

and pubescent towards the base. Staminal sheath

curved, ca. 17 mm long, glabrous. Ovary shortly

stipitate, hairy, with ca. 12 ovules; style slender,

flattened, glabrous except near the base. Pods not

known.—Fig. 2A-G.

Distribu tion. —Only known from the lower

Mangoro River valley in E Madagascar.

HABITAT.—In remnants of humid, eastern,

lowland forest, at ca. 200 m altitude.

FLOWERING TIME.—Only recorded in October.

Acknowledgements

We thank A. Beaumont and D. Storez for the

illustrations, and J.-J. FLORET for assistance with the

Latin diagnoses. D.J. Du PüYwould like to thank the

Royal Society for the opportunity to undertakc colla¬

borative research in the Laboratoire de Phanérogamie,

Paris, and the Weston Foundation for the support of

his research in Madagascar and Kew. We would also

like to thank the Directors and staff of the Laboratoire

de Phanérogamie, Paris, the Royal Botanic Gardens,

Kew, the Parc de Tsimbazaza, Antananarivo, and the

Centre National de la Recherche sur l’Environ¬

nement, Antananarivo.

REFERENCES

Du Puy D.J. & Labat J.-N. 1995.— Pyrantbus , a new

genus of the tribe Millettieae (Leguminosae-

Papilionoideae). Kew Bull. 50: 73-84.

Du Puy D.J., Labat J.-N. & Schruie B.D. 1994.—

Révision du genre Vaugbania S. Moore

(Leguminosae-Papilionoidcac-Indigofereae). Bull.

Mus. Natl. Hist. Nat., B, Adansonia 16: 75-102.

Du Puy D.J., Labat J.-N. tk Schrire B.D. 1995.— A

révision of Phylloxylon (Leguminosae-

Papilionoideae: Indigofereae). Kew Bull. 50: 477-

494.

GEESINK R. 1981.—Tephrosieae: 245-260, in

POLHIEL R.M. & Raven P.H. (eds.), Advances in

Legutne Systematics 1. Royal Botanic Gardens, Kew.

GeESINK R. 1984.— Scala Millctuatrum A stervey of

the généra of the tribe Millettieae ( l.egum.-Pap.) with

methodological considérations. E.J, Brill/Leiden

University Press, Leiden Botanical Sériés 8.

HUANC S.-F. 1991.'— Strongylodon (Leguminosae-

Erythrininae), a révision of the genus. Wageningen

Agric. Univ. Papers 90: 1-69.

LABAT J.-N. 1996.—Biogéographie, endémisme et

origine des Leguminoseae-Papilionoideae de

Madagascar: 95-108, in LOURENÇO W.R. (ed.),

Biogéographie de Madagascar. ORSTOM, Paris.

Labat J.-N. & Du Puy D.J. 1995.—New species and

combinations in Millenia Wight & Arnott and

Pongamiopsis R. Viguicr (Legliminosae-

Papilionoideae-Millettieae) from Madagascar.

Novon 5: 171-182.

POLHILL R.M. 1981.—Papilionoideae: 191-208, in

Polhill R.M. & Raven P.H. (eds.), Advances in

Legtime Systematics 1. Royal Boranic Gardens, Kew.

SCHRIRE B.D. 1995.—Evolution of the Tribe

Indigofereae (Leguminosae-Papilionoideae): 161-

244, in Crisp M.D. & Doyle J.J. (eds.), Advances

in Legutne Systematics 7, Phylogeny. Royal Botanic

Gardens, Kew.

VlGUiER R. 1952.—Leguminosae madagascarienses

novae (suite 2). NotuL Syst. (Paris) 14: 168-187.

Manuscript received 15 January 1998;

revised version accepted 13 March 1998.

ADANSONIA, sér. 3 • 1998 • 20(1)

171

Impact des feux de brousse sur les maquis

ligno-herbacés des roches ultramafiques

de Nouvelle-Calédonie

Tanguy JAFFRÉ, Frédéric RIGAULT & Gilles DAGOSTINI

Laboratoire de Botanique et d’Écologie Végétale,

Centre ORSTOM, BP A5, 98848 Nouméa, Nouvelle-Calédonie.

jaffre@noumea.orstom.nc

MOTS CLÉS

incendies,

maquis,

roches ultramafiques,

flore endémique,

Cyperaceae,

autosuccession,

Nouvelle-Calédonie.

RÉSUMÉ

L’effet des feux de brousse sur deux catégories de maquis ligno-herbacés, se

développant sur sols issus de roches ultramafiques, a été suivi sur une période

de plus de 10 ans, par la méthode des points quadrats. Au terme des observa¬

tions, on assiste à la reconstitution presque complète du cortège floristique

initial. La majorité des espèces arbustives se régénèrent par rejets. Toutefois

cinq espèces ne se réinstallent que très progressivement, par germinations de

semences. Le changement le plus important est d’ordre structural et inter¬

vient au niveau de la strate herbacée. Les Cyperaceae cespiteuses, entièrement

détruites par l’incendie, ne se réinstallent que très lentement à partir de

semences. L’espace laissé libre est largement occupé par une espèce joncifor-

me rhizomatcuse (Lepidosperma perteres) qui, contrairement aux Cyperaceae

précédentes, s oppose à la progression du feu. La reconstitution du tapis

végétal après incendie s’apparente donc a une « autosuccession » qui permet

au maquis, si aucun facteur n'intervient pour provoquer un nouvel incendie,

de retrouver à terme ses caractéristiques initiales. Toutefois, l’interruption du

processus de reconstitution par un incendie précoce, entraîne une proliféra¬

tion de Pteridium esculentum. Cette espèce, très inflammable, favorise la pro¬

pagation du feu dont les effets répétés se traduisent par l’installation d’une

fougeraie permanente.

ADANSONIA, sér. 3 • 1998 • 20(1)

173

Jaffré T., Rigault F. & Dagostini G.

KEYWORDS

fire,

maquis,

ultramafk rock,

endemic flora,

Cyperaccac,

autosuccession,

New Caledonia.

ABSTRACT

The effect of fire on rwo types of woody-herbaceous maquis on soils derived

form uktamafic rocks was examined for over ten years using the line transect

method. During the course of the study, the initial floristic composition was

almost completely restored. Most of the woody species resprout, whereas five

fire sensitive species reestablish slowly on hurnt areas front seed. The most

important change was structural, taking place in the herbaceous layer.

Tussock Cyperaeeac, emirely elitninated by the fire, regtew very slowly from

seed. Open areas were largely occupicd by a single rhizomatous species

(Lepidospcnnii perteres) whiclt, unlike the Cyperaeeac présent beforc burning,

is résistant to the spread of fire. The formation of a new plant cover after

burning thus represents “atitosuccessioti", CHabling the maquis to return to

its initial charactcristics provided thaï ntt additional fires occur. This process

of végétation recovery can however bc interrupted by prématuré burning,

vvhich brings about a prolifération of Pteriaium esculentum, a highly fiam-

mable species that facilitâtes the spread of fire. Rcpeated burning can thus

resuit in the establishment of a permanent fern cover.

INTRODUCTION

Les feux de brousse constituent dans l’ensemble

du monde tropical et méditerranéen, comme en

témoignent de nombreux ouvrages (GlLL et al.

1981 ; Booysfn & Talnton 1984 ;

GOLDAMMFK 1990 ; VAN WlLGEN et al. 1992 ;

WHELAN 1995), l'un des facteurs principaux de

dégradation et de transformation du milieu natu¬

rel. Ceci est également le cas, en Nouvelle-

Calédonie, où, comme souligné par VlROT

(1956) les feux de brousse ont eu un rôle impor¬

tant dans la configuration et la distribution

actuelles des formations végétales.

On peut à ce jour estimer que la totalité des

savanes, des fourrés et des maquis sur roches

acides, soit un total de 6500 km 2 , en sont le

résultat. Ces formations, dont la flore est compo¬

sée majoritairement d’espèces introduites, ont été

classées par MORAT et al (1981) en « formations

transformées ». Elles s’opposent aux « formations

autochtones » dont la flore est très largement

constituée d’espèces endémiques de la Nouvelle-

Calédonie, Ceci n’exclut pas que certaines forma¬

tions autochtones de type édaphique comme les

« maquis miniers » résultent pour la plupart de la

destruction du couvert forestier initial (Jaffré

1980 ; Morat et al. 1986). — Fig. 1.

Bien que l’on air tendance à considérer que les

feux ont débuté avec l’arrivée de l’homme, esti¬

mée à environ 4000 BP pour la Nouvelle-

Calédonie, des études récentes (HOPF., sous

presse), basées sur l'examen de pollens fossiles

recueillis dans des sédiments anciens de la Plaine

des Lacs, remettent en cause ce point de vue. Les

résultats montrent l’alternance depuis 30.000 ans

de phases forestières et de phases pionnières

« posr incendie » marquées localement par la

dominance d'espèces pionnières du genre

Gymnastoma (Casuarinaceae). L’adaptation au

feu des espèces néo-calédoniennes pourrait donc

être beaucoup plus antienne que ce que l’on a

coutume de considérer.

La gravité des atteintes provoquées par des feux

de brousse au couvert végétal actuel, est large¬

ment fonction des caractéristiques des groupe¬

ments végétaux touchés (JAFFRÉ et al. 1997).

Aussi une meilleure connaissance des effets

immédiats et à plus long terme, des incendies sur

les différentes catégories de végétation, est-elle

nécessaire pour définir les modalités de gestion,

adaptées à chaque cas.

174

ADANSONIA, séf. 3 • 1998 • 20(1)

Feux en maquis ligno-herbacés de Nouvelle-Calédonie

Si la foudre peut être à l'origine de quelques

foyers, il est certain que les incendies se sont

intensifiés depuis J’arrivée des premiers hommes

en Nouvelle-Calédonie (estimée à 4000 BP) et

sont largement liés aux pratiques culturales. Les

feux sont utilisés comme auxiliaires nécessaires

au défrichement des terres destinées à l’agricultu¬

re et, depuis l'arrivée des Européens, à I élevage.

Ils se sont accrus aux cours des dernières décen¬

nies avec le développement économique du

Territoire et l’ouverture de nouvelles voies

d’accès. Dans les secteurs sans valeur agricole, les

feux sont associés à des pratiques de chasse et à

des prospections minières et aussi, bien souvent,

à la propagation accidentelle non contrôlée des

feux allumés initialement pour défricher des

zones à vocation agro-pastorale. Toutefois beau¬

coup d’incendies, qui n’ont pas de causes identi¬

fiées, sont dus à des négligences et à des actes de

malveillance.

CADRE ET MÉTHODES D’ÉTUDE

Cette étude concerne l’effet des incendies sur

une catégorie de « maquis minier », le « maquis

ligne herbacé » et a pour but, d’une part d’éva¬

luer la nature, l’importance et la durée des modi¬

fications du couvert végétal, d’autre part

d’analyser le processus de sa reconstitution.

Le terme de « maquis minier » dans son accep¬

tion locale « regroupe toutes les formations sur

roches ultramafiques (péridotites et serpentïnites)

n’appartenant pas à la forêt dense humide ou aux

forêts rivulaires. Il regroupe des formations sclé-

tophylles sempervirentes héliophiles, arbustives

plus ou moins buissonnantes ou ligno-herbacées

à strate cypéraoécnne dense » (MORAT et al.

1986). Ces dernières, qui sont les seules concer¬

nées par la présente étude, occupent des zones

anciennement boisées. Leur flore est constituée,

comme celle de l’ensemble des maquis miniers,

20 °-

0 50 km

Végétation intacte ou peu modifiée

Hl Forêts denses humides

|g| Forêts sclérophylles

| | Maquis miniers

Végétation modifiée

□ Savanes et fourrés divers

EHH Maquis miniers secondaires

164°E

Fig. 1. — Répartition des principales formations végétales de la Grande-Terre (Nouvelle-Calédonie), et localisation du secteur de la

Plaine des Lacs.

ADANSONIA, sér. 3 • 1998 • 20(1)

175

Jaffré T., Rigault F. & Dagostini G.

par un fort pourcentage (entre 85 et 90%)

d’espèces endémiques du Territoire (JAFFRÉ et al.

1994).

Les maquis ligno-herbacés sur roches ultrama-

fiques se distinguent des autres catégories de

maquis par une strate herbacée très développée

(recouvrement > 75%). Celle ci est formée en

majorité de Cyperaceae cespiteuses appartenant

aux genres Costulariu et Schoenus, très inflam¬

mables en période sèche et d’une espèce joncifor-

me plus discrète, Lepidospernw perteres. Le taux

de recouvrement de la strate herbacée est supé¬

rieur à celui de la strate arbustive qui peut être

buissonnante et discontinue d’une hauteur com¬

prise entre 20 cm er 1,3 m.

Ce type de maquis se trouve sur sols oxydiques

(ferrallitiques ferridques) remaniés par érosion et

colluvionnement sur les versants ou en situation

de piémont à des altitudes variées, du niveau de

la mer à 1600 m d'altitude. II ne se trouve pas

sur les sols oxydiques cuirassés ou gravillonnaires

peu propices au développement des Cyperaceae.

L’effet des incendies sur les maquis occupant ces

derniers substrats a été étudié séparément (Mac

Coy et al., sous presse ; RlGG et al., sous presse).

L'étude a débuté en octobre 1984, trois

semaines après le passage du feu. Elle porte sur

un maquis de piémont et un maquis de haut ver¬

sant de la Plaine des Lacs (Fig. I) situés entre

230 et 260 m d’altitude, à l’extrême sud de la

Grande-l’erre. La pluviométrie annuelle moyen¬

ne de la zone s'établit entre 2500 et 3000 mm.

Les mois les plus secs, avec des précipitations

inférieures il 100 mm, se situent de manière assez

irrégulière de septembre à novembre. La tempé¬

rature annuelle moyenne est de l’ordre de 21°, le

mois le plus chaud étant février (25°) et le mois

le plus frais, août ( 18").

En se référant aux travaux antérieurs réalisés

dans Je Grand Massif du Sud (JAFFRF 1980) il est

possible de rattacher le maquis de haut versant à

l’association à Coslularia pubescens et Stypbelia

albicans , le maquis de piémont à l’association à

Codia discolor et Eugénie! strie ta. Dans les deux

cas, il s'agit d’une végétation peu dynamique

constituée d’espèces adaptées à des conditions de

nutrition minérale extrêmes en raison de la pau¬

vreté du sol en P, K et Ca et à des teneurs anor-

malement élevées en nickel, chrome et

manganèse (Tableau 1).

Tableau 1. — Composition chimique moyenne de l'horizon supérieur (2-10 cm) des sols de hauts versants et des

sols de piémonts.

Stations

Analyses

Haut versant

Piémont

6,2 ±1,4

5,4 ±0,5

Bases échangeables

(me /100 g)

Ca ++

0,85 ±0,53

0,37 ±0,34

Mg ++

1,54 ± 1,02

0,46 ± 0,38

K +

0,05 ±0,02

0,05 ± 0,03

Na +

0,07 ±0,02

0,05 ± 0,03

Éléments totaux

P ppm

227±175

241 ±143

Ca%

0,01 ±0,01

0,01 ±0,00

Mg %

0,65 ± 0,38

0.33 ± 0,28

K %

0,02 ±0,02

Na%

0,04 ± 0,04

0,04 ±0,06

Mn%

0,45 ±0,17

0.63 ± 0,37

Ni %

0,96 ±0,21

0,68 ±0,22

Cr %

2,5 ±1,5

2.4 ±1,1

Fe %

41,7 ±8,0

46,1 ±3,8

176

ADANSONIA, sér. 3 • 1998 • 20(1)

Feux en maquis ligno-herbacés de Nouvelle-Calédonie

En raison de l'importance des Cyperaceae cespi-

teuses et stolonifères pour lesquelles la notion

d’individu est souvent ambiguë, l'évolution du

tapis végétal a été suivie par la méthode linéaire

des points quadrats (CanFIEI I) 1941) dont la

technique a été reprise et décrite pour une étude

similaire par FORGPAUD & TotJI-HT (1979).

Chaque ligne élémentaire a été matérialisée sur

le terrain par 2 piquets entre lesquels est tendue

une cordelette à laquelle est superposé un ruban

métallique gradué. Les lectures ont été effectuées

tous les 10 cm à différentes périodes, de la date

de I incendie il 1997 pour la zone de haut versant

et seulement jusqu'en 1995 pour la zone de pié-

mont, qui en 1996 à été défrichée pour une opé¬

ration de reboisement.

Le principe de la méthode consiste à noter les

espèces touchant une tige métallique de faible

diamètre plantée perpendiculairement au sol. Les

données recueillies permettent de calculer à un

temps donné :

— le recouvrement végétal, qui correspond au

pourcentage de points où une ou plusieurs

espèces sont en contact avec la tige de mesure ;

— la fréquence relative des diverses espèces,

représentée par le pourcentage de points de lectu¬

re où l'espèce est notée ;

— la contribution spécifique des diverses

espèces, qui est le rapport de la fréquence relative

d’une espèce à la somme des fréquences relatives

de toutes les espèces.

En outre, toutes les espèces ont été inventoriées

sur 2501) nr de la zone brûlée et des plants des

différentes espèces ont été déterrés afin de déter¬

miner leurs modes de régénération et de multi¬

plication.

Dans chacune des deux zones, les lignes perma¬

nentes ont été établies au sein de surfaces homo¬

gènes représentatives de l'ensemble ; elles

totalisent 170 m (6 lignes de 25 m et une ligne

de 20 m) en position de haut versant et 150 m (6

lignes de 25 m) en zone de piémonr.

RÉSULTATS

Évolution de la composition et de la richesse

floristique

La comparaison de la composition floristique

initiale et de celle observée au terme de 13 ans

pour le haut versant et de 11 ans pour le piémont

montre une grande stabilité. En effet, 44 espèces

sur les 46 initiales se retrouvent dans la parcelle

de haut versant et 40 sur 41 dans celle de pié¬

mont (Tableau 2).

On note seulement la disparition de

Myodocarpus lanceolatus (Araliaceae) dans les

deux stations et celle de Styphelia albicans

(Epacridaceae) dans le maquis de haut versant.

L’absence de Myodocarpus lanceolatus s’explique

par son caractère pré-forestier, commun à plu¬

sieurs espèces de la famille des Araliaceae de la

flore de Nouvelle-Calédonie. L'absence de

Styphelia albicans tient à la lenteur de la plupart

des espèces du maquis minier à se réinstaller à

partir de semences. Ainsi Styphelia cymbnlae et

Dracophyllum rarnosum, deux autres espèces de la

famille des Epacridaceae qui ne rejertent pas de

souche, n'ont été observées respectivement sur les

lignes de lecture que 5 et 7 années après l’incen¬

die.

Les espèces supplémentaires ne sont qu’au

nombre de deux, Baumea deplanchei

(Cyperaceae) (espèce pionnière des surfaces nues,

endémique des sols issus de roches ultrama-

ftqties), commune aux deux stations, et f'inus

caribaea (espèce introduite, plantée dans le sec¬

teur en 1965) dont trois individus se sont instal¬

lés dans la parcelle de piémont, où il montre

toutefois un développement limité L’observation

de maquis miniers n'ayant pas brûlé depuis plu¬

sieurs dizaines d’années permet de constater que

Baumea deplanchei ne persiste pas lorsque la stra¬

te herbacée, a base de Cyperaceae cespiteuses, est

redevenue dense Par contre P inns caribaea

demeure er devient parfois envahissant, notam¬

ment dans les zones de piémont à proximité des

plantations.

Evolution du taux de recouvrement végétal

L’évolution du taux de recouvrement de la végé¬

tation, dans chacune des stations étudiées, est

représentée sur la Figure 2.

On observe dans les deux cas une reprise rapide

de la végétation qui, d'une manière générale,

débure dès la première pluie après l’incendie.

Bien que le haut versant soit l’objet d’une érosion

consécutive au feu, son recouvrement végétal

ADANSONIA, sér. 3 • 1998 • 20(1)

177

Jaffré T., Rigault F. & Dagostini G.

% recouvrement

8 â S

maquis de haut versant

> M • * maquis de piémont

Fig. 2. — Evolution du taux de recouvrement de la végétation des deux stations étudiées.

demeure supérieur à celui du piémont. Les taux

de recouvrement croissent rapidement au cours

des deux premières années et marquent un flé¬

chissement à parrir de la quatrième année. Sur

haut versant, les zones dénudées correspondent à

des blocs rocheux et à de petites ravines d’érosion

qui se creusent à chaque forte pluie. Sur piémont

ce sont des plages de gravillons, inondées en

période de pluies abondantes et arides en période

sèche Les taux de recouvrement mesurés au

terme des observations (environ 75% sur haut

versant et 60% sur piémont) sont assez proches

de ceux généralement observés pour les maquis

ligno-herbacés voisins, épargnés par le feu depuis

plusieurs décennies. Toutefois la végétation

demeure dans les deux zones étudiées, moins

haute et moins dense en raison d’une faillie

superposition des frondaisons des différentes

espèces.

Evolution des composantes de la végétation

En fonction de leur morphologie et de leur

dynamique, on peut distinguer 5 composantes de

la végétation :

— Pteridium esculentnm, fougère rhizomateuse,

très discrète et souvent absente des maquis épar¬

gnés par le leu depuis plusieurs décennies, qui

émet de nombreuses frondes après l’incendie ;

— Lepidosperma perteres , Cyperaceae, espèce

jonciforme rhizomateuse, très clairsemée dans la

végétation initiale, qui émet d’abondantes tiges

après incendie ;

— Lensemble des Cyperaceae cespiteuses, qui

constituaient la plus grande partie du couvert

herbacé (recouvrement de l’ordre de 75%) et de

la biomasse facilement combustible avant

l'incendie.

— L’ensemble des espèces arbustives, dont le

recouvrement était compris entre 30 et 50%

avant l’incendie ;

— L’ensemble des espèces n’appartenant pas

aux catégories précédentes : Ditmella stipitata

(Liliaceae), Eriaxis rigida et Megastylis gigas

(Orchidaceae), Schizaea dichotoma et S, laevigata

(Schizaeaceae), dont la somme des importances

relatives n’excède à aucun moment 2,75% et

dont les variations sont, compte tenu des marges

d’erreur, trop faibles pour être interprétées.

178

ADANSONIA. sèr. 3 • 1998 ■ 20(1)

Feux en maquis ligno-herbacés de Nouvelle-Calédonie

% fréquence relative

(Q O •=.

— — Pteridlum esculentum

_ ^ — Cyperaceae cespiteuses

* -

Lepidosperma perteres

Ensemble des espèces arbustives

Fig. 3. — Évolution de la fréquence relative des quatre principales composantes de la végétation sur haut versant.

% fréquence relative

•=. ro

— ^ — Pteridium esculentum

Lepidosperma perteres

^ _ Cyperaceae cespiteuses ■ ■ ^ • * Ensemble des espèces arbustives

Fig. 4. — Évolution de la fréquence relative des quatre principales composantes de la végétation sur piémont.

ADANSONIA. sér. 3 • 1998 • 20(1)

Jaffré T., Rigault F. & Dagostini G.

% contribution spécifique

70 -

Maquis de haut versant

v ~*-- «.

.X-

r- - *

oct. 84

avr. 85

août 86

Juin 87

avr. 88

oct. 88

juin 89

nov. 89

avr. 92

déc. 93

août 95

— — Pteridium esculentum

-•-

Lepidosperma perteres

— _ Cyperaceae cespiteuses

Ensemble des espèces arbustives

Fig. 5. — Évolution de la contribution spécifique des quatre principales composantes de la végétation sur haut versant.

% contribution spécifique

70 T

Maquis de piémont

Pteridium esculentum

Cyperaceae cespiteuses

Lepidosperma perteres

_ Cyperaceae cespiteuses Ensemble des espèces arbustives

Évolution de la contribution spécifique des quatre principales composantes de la végétation sur piémont.

ADANSONIA, sér 3 • 1998 • 20(1)

Feux en maquis ligno-herbacés de Nouvelle-Calédonie

L’évolution de la fréquence relative des quatre

principales composantes de la végétation est

représentée sur la Figure 3 pour la station de

haut versant et sur la Figure 4 pour celle de pié-

mont. L’évolution de la contribution spécifique

est pour sa part représentée sur la Figure 5 pour

le maquis de haut versant et sur la Figure 6 pour

celui de piémont.

— Evolution de Pteridium esculentum.

Dans chacune des stations, l’installation de

cette espèce est immédiate, puisqu’elle émet de

nouvelles frondes très rapidement après l’incen¬

die. Ses fréquences relatives, de 15% sur haut

versant et d’environ 10% sur piémont, données

dès la première lecture, demeurent relativement

stables au cours des 4 à 5 premières années

d’observation. Elle décroît ensuite sensiblement

pour atteindre finalement des valeurs de 3% sur

haut versant après 13 ans et de 5% sur piémont

au bout de 11 ans.

Pteridium esculentum présente la contribution

spécifique la plus importante 3 semaines après

l’incendie. Elle s’établit à 44% sur piémont et

65% sur haut versant. Mais elle décroît ensuite

rapidement, surtout les deuxième et troisième

années et atteint finalement dans les deux cas,

une valeur en dessous de 10% au bout d’une

dizaine d’années, soir la valeur la plus basse des 4

composantes principales du couvert végétal.

— Evolution de Lepidosperma perteres.

La fréquence relative de cette espèce joncifor¬

me, qui émet d’abondantes tiges dès les pre¬

mières semaines suivant l'incendie, ne cesse de

croître pendant les 5 à 8 premières années pour

atteindre respectivement 45% et 50% sur pié¬

mont et sur haut versant. Au cours des années

suivantes on note un fléchissement de la courbe

qui atteint finalement dans les deux cas une

valeur voisine de 40%.

Lepidosperma perteres possède, dès les premiers

mois sur piémont et au bout de moins de 2 ans

sur haut versant, une contribution spécifique

supérieure à celle des autres composantes de la

végétation. Sur haut versant la valeur de la

contribution spécifique reste relativement stable

entre 2 er 5 ans puis décroit progressivement, à

partir de la cinquième année. Sur piémont cette

décroissance est sensible dès la quatrième année.

Dans chacun des cas elle atteint, au terme des

observations, des valeurs voisines de 40%. A ce

stade, la contribution spécifique de Lepidosperma

perteres est égale ou supérieure à celle des autres

composantes de la végétation mais se trouve dans

une phase décroissante alors que la contribution

spécifique des autres composantes est dans une

phase croissante.

La prépondérance de Lepidosperma perteres , qui

occupe l’espace laissé libre par la destruction des

Cyperaceac cespireuses, donne aux deux maquis

étudiés une physionomie particulière (« maquis

jonciforme ») qui, sur le terrain, permet aisément

d’identifier les zones ayant brûlé il y a 5 à 10 ans.

— Evolution de l’ensemble des espèces arbus-

tives.

La fréquence relative des espèces arbustives,

dont le plus grand nombre rejette de souche

après l incendie, plus ou moins rapidement selon

les conditions hydriques liées à la pluviométrie,

croît assez régulièrement au cours des périodes

d’observation. Elle atteint 23% en zone de pié¬

mont et 35% sur haut versant au bout, respecti¬

vement, de 11 et 13 ans.

La contribution spécifique des espèces arbus¬

tives croît lentement dans les deux stations. Elle

atteint 24% en zone de piémont et 28% en posi¬

tion de haut versant après 11 années et 35% sur

haut versant après 13 ans. Dans ce dernier cas, la

contribution spécifique des espèces arbustives, en

phase croissante, est égale à celle de Lepidospenna

perteres.

— Évolution de l’ensemble des Cyperaceae ces-

piteuses.

La strate de Cyperaceae cespiteuses ( Costularia,

Schoenus), la plus vulnérable au feu, ne subsiste

après l’incendie qu’à l’état de touffes isolées loca¬

lement épargnées par les flammes. Sa réinstalla¬

tion s’effectue très lentement par semences au

cours des 4 à 6 premières années suivant Lincen¬

die et s’amplifie ensuite, de manière plus nette

sur piémont que sur haut versant. Au terme des

observations sa fréquence relative, en phase crois¬

sante, atteint 30% sur piémont et 25% sur haut

versant.

La contribution spécifique des Cyperaceae ces-

pitcuses demeure inférieure à celle des 3 autres

principales composantes de la végétation jusqu’à

la cinquième année sur piémont et jusqu’à la hui¬

tième année sur haut versant, atteignant dans

ADANSONIA. sèr. 3 • 1998 • 20(1)

181

Jaffré T., Rigault F. & Dagostini G.

chaque cas une valeur de l'ordre de 10%.

Ensuite, elle s’accroît plus rapidement sur pié-

mont que sur haut versant. Elle atteint dans le

premier cas une valeur de 28% au bout de 11 ans

et se situe en deuxième position juste en dessous

de LepidospermU pertem. Dans le second cas elle

atteint 18% la onzième année et 25% au bout de

13 ans. À ce stade elle se trouve en troisième

position, en dessous de Lepidospenna pertem et

de l’ensemble des espèces arbustives.

Réaction des espèces à l’action du feu

Les modes de réinstallation des différentes

espèces observées dans les deux stations sont

donnés dans le Tableau 2.

Tableau 2.—

Peu d'espèces meurent massivement après

l’incendie. C'est toutefois le cas, comme il a été

noté plus haut, des Cyperaceae cespiteuses,

notamment de Costularia nervosa et à un degré

moindre de Costularia comosa, C.. stagnait), C.

antndittacea , Schoenus neocaledonicus et Sch. juve-

nis. C.’est aussi le cas de quelques espèces arbus¬

tives : Dracnphyllmn ramusum . Styphelia cymhulae

et 5. albicans (F.pacridaceae), Myodocarptts lanceo-

latus (Araliaceae) et Alphitonia tieocaledonica

(Rhamnaceae). Ces différentes espèces ne se

réinstallent ensuite qu'à partir de graines. Dans le

cas des Cyperaceae comme dans celui

d Alphitonia neocaledonica , la régénération s’effec¬

tue à partir de semences contenues dans le sol

Composition floristique et modes de réinstallation des espèces des deux parcelles étudiées.

Espèces

Piémont

Haut versant

Mode de

réinstallation

Agavaceae

Cordyline neocaledonica Linden

Apocynaceae

Melodinus balansae Baill.

Parsonsia populifolia Baill.

Araliaceae

♦ Myodocarpus lanceolatus Dubard & R. Vig.

Asclcpiadaccae

Marsdema microstoma Schltr.

Celastraceae

PenptnryQni margmata (Baill.) Loes.

Cunonlaceae

Codia dir.coloi (Biongn. & Gris) Guillaumin

Codia mtida Schltr

Pancheria alalernoides Brongn.

Pancheria hirsula Vieill. ex Pamp.

Pancheria vigillardii Brongn.

Cyperaceae

• Baumoa deplanchci Boeck.

Costulaiia aivndinacea (Sol. ex Vahl) Kük.

Coslularia mmosa (CB. Clarke) Kük.

Costularla nervosa J. Raynal

Coslularia pubescens J. Raynal

Costularia stagnalis (Dâniker) Kük.

Lepldosperma perteres C.B. Clarke

Schoenus jtivcnis C.B. Clarke

Schoenus neocaledonicus C.B Clarke

Dennstaedtiaceae

Ptoridium osculenlum (Forster f.) Cockayne

Dllleniaceae

Hibbetlia pancheri (Brongn. & Gris) Briq.

H/bbertia pulchella (Brongn. & Gris) Schltr.

Hlbbertta Irachypbylla Schltr.

Epacridaeeae

♦ Styphelia albicans (Brongn. & Gris) Sleumer

182

ADANSONIA, sér. 3 • 1998 • 20(1)

Feux en maquis ligno-herbacés de Nouvelle-Calédonie

Styphelia cymbulae (Labill.) Sprengel

Dracophyllum ramosum Pancher ex Brongn. & Gris

Escalloniaceae

Argophyllum montanum Schltr.

Eupnorbiaceae

Phyllanlhus aeneus Baill.

Phyllanlhus chrysanthus Baill.

Flacourtiaceae

Casearia silvana Sch Itr.

Goodenfaceae

Scaevoia beckn Zarilbr.

Guttiferae

Garcinia amplexicaulis Vieill.

Monlrouziera sphaeroidea Pancher ex Planch. & Triana

Liliaceae

Dianalla stipitata Schlittler

Linaceae

Hugonia penicillanthemum Baill, ex Pancher & Sebert

Loganiaceae

Geniostama celaslrineum Baill.

Moraceae

Ficus aspera G Forst.

Myrtaceae

Austromyrtus alaternoides (Brongn. & Gris) Burret

Babmqtonia leratii (Schltr.) A.R. Beans

Cloezia artensis (Monfr.) P S. Green

Eugenla stricts Pancher ex Brongn. & Gris

Tristaniopsis glauca Brongn. S Gris

Uromyrtus emargmata (Pancher ex Brongn. & Gris) Burret

Xanthoslemon aurantiacum (Brongn. & Gris) Schltr.

Orchidaceae

Ertaxis rigida Reichb. f

Megastylis gigas (Reichb. f.) Schtr.

Pinaceae

• Pinus caiibaea Morelet

Praleaceae

Greyillea eXUI Lindley subsp. rubiginosa (Brongn. & Gris) Virot

Grevillea gillivrayi Hook. & Arn.

Stonoc<vpus umoeihferus (J R Forst.) Druce

Rhamnaceae

Alphifonia naocaledonica (Schltr.) Guillaumin

Rubtaceae

Normandia neocaledonica Hook. f.

Psychotria olnoides (Baill.) Schltr.

Sapïndaceae

Guioa glauca (Labill.) Radlk.

Guioa villosa Radlk.

Storthocalyx pancheri (Baill.) Radlk.

Schîzaeaceae

Scbizaea dichotoma (L.) Smith

Schizaea laevlgata Mett.

Smilacaceae

Srwlaxsp. (Vetllon 3882)

Thymeliaceae

Wikslroemia indica (L.) C. Meyer

Violaceae

Agalea deplanchei Brongn. & Gris ex Guillaumin

• espèce n'appartenant pas à la flore initiale R rejets

♦ espèce non répertoriée au terme des observations S semences

ADANSONIA. sér. 3 • 1998 • 20(1)

183

Jaffré T., Rigault F. & Dagostini G.

qui germent dès les premières pluies après

l’incendie. Les autres espèces, comprenant toutes

les Epacridaceae, ne se réinstallent que plus tard

par apports successifs de semences.

La majorité des espèces des rnaquis-ligno-herb3-

cés résistent au feu, bien que la partie aérienne

soit entièrement détruite. C’est le cas parmi les

espèces herbacées, des Orchidaceac, Megastylis

gigas et Eriaxis rigida, qui fleurissent abondam¬

ment dès les premières pluies suivant l’incendie,

ainsi que celui des espèces rhizomateuses,

Pteridium escukntum, Schiztlea spp. (fougères),

Lepidospermu perteres (Cyperaceae), Diavelln sti-

pitata (Liliaceae). C’est aussi le cas de la majorité

des espèces arbustivcs qui, après l incendie, déve¬

loppent des rejets à croissance rapide, à partir de

bourrelets ligneux (lignotuber) situés au niveau

du collet (Fig. 7, 8). Elles appartiennent à des

familles variées (Myrtaceae, Cunoniaceae,

Dilleniaceae) dont les espèces sont souvent domi¬

nantes dans la strate arbustive des maquis

miniers.

Ces bourrelets ligneux et robustes peuvent

atteindre jusqu’à 1 m de diamètre pour certains

Tristaniopsis spp. (Myrtaceae), témoignant de la

longévité de ces espèces et de l’ancienneté des

incendies répétés.

Quelques espèces, dont Guioa villosa

(Sapindaceae) et Montrouziera sphaeroidea

(Guttiferae) émettent des rejets de racines après

l’incendie. Parmi les espèces ligneuses résistantes

au feu, une seule, Babingtonia leratii (Myrtaceae)

se régénère dès la première année suivant l’incen¬

die, simultanément à partir de rejets et de germi¬

nations.

DISCUSSION ET CONCLUSION

Au stade actuel des observations, on constate

que l’effet de l’incendie se manifeste de manière

sensiblement identique sur les deux catégories de

maquis ligno-herbacés. Les seules différences

étant le développement plus important de la stra¬

te arbustive sur haut versant que sur piémont,

l’inverse s’observant pour la strate des Cyperaceae

cespiteuses.

Les suivis réalisés sur ces deux stations pendant

plus de 10 ans, complétés par plusieurs autres

observations sur les maquis ligno-herbacés du

même secteur géographique, permettent de faire

un certain nombre de commentaires et de tirer

quelques conclusions.

Les modifications de la composition floristique

provoquées par l’incendie sont peu importantes

et ne semblent pas définitives. En effet, les

quelques espèces qui ne sont pas réapparues,

demeurent largement représentées dans des

maquis voisins n’ayant pas brûlé depuis au moins

30 années. En outre, parmi les espèces supplé¬

mentaires, Baume a depbinchei , dont la présence

demeure discrète dans les deux cas étudiés, et qui

est absente des maquis plus anciens, ne s installe

manifestement pas de manière durable. Par

contre, Pin us earibaea qui s implante volontiers

sur les piémonts, peut s’y maintenir et sans

doute, à plus long terme, s’y multiplier. On n’a

pas encore pour cette espèce (introduite dans les

années soixante) suffisamment de recul pour tirer

des conclusions définitives sur le rôle du feu dans

sa prolifération.

D’une manière générale, l’effet de l’incendie se

traduit principalement par un changement de la

structure du maquis, engendré par la modifica¬

tion ife l’abondance relative des principales com¬

posantes de la flore. Ainsi assisre-t-on au

remplacement de la strate cypéracéenne cespiteu-

se par une strate cypéracéenne jonciforme- Ce

changement s’accompagne du développement,

variahle suivant les sites, de Pteridium esculentum,

qui toutefois périclite progressivement dans les

conditions habituelles, comme cela a été montré

dans les deux stations étudiées. L’incendie a aussi

pour effet de détruire la totalité de la biomasse

ligneuse aérienne et par contrecoup, d'induire

l’émission de jeunes pousses chez la plupart des

espèces arbustives, sauf chez quelques-unes

appartenant notamment aux familles des

Epacridaceae et des Araliaceae. Ces dernières se

réinstalleront toutefois progressivement à partir

de semences. Aussi, malgré une émission rapide

de rejets, la strate arbustive mettra plus de 10

années pour se reconstituer.

La strate cypéracéenne cespiteuse, qui constitue

un matériel végétal très inflammable en saison

sèche, ne se réinstalle que très lentement à partir

de semences et laisse place à une strate cypéra¬

céenne jonciforme ininflammable qui met le

186

ADANSONIA.sér. 3 • 1998 • 20(1)

Feux en maquis ligno-herbacés de Nouvelle-Calédonie

maquis à l’abri des feux pendant plusieurs

années. Ceci à condition toutefois qu’un déve¬

loppement trop important de Pteridium esculen-

tum n’entraîne pas une accumulation de frondes

sèches, propice à la propagation des incendies.

La succession secondaire après incendie des

maquis ligno-herbacés, qui se limite donc à un

phénomène de retour au stade initial, sans passa¬

ge par des stades successifs intermédiaires, floris-

tiquement différenciés, est très largement

déterminée par la composition floristique avant

l’incendie. Le phénomène s’apparente donc au

modèle de la composition initiale proposé par

EGLER (1954).

Ce phénomène, qui se caractérise finalement

par l'absence de changement de la composition

floristique est assez général ; il a été décrit dans le

Bassin Méditerranéen (TrabauT 1987 ; CODV &

MoONEY 1976), en Australie (RUSSELL &

Parson 1978 ; Specht, Rayson & Jackman

1958), en Afrique du Sud (VAN WlLGEN 1981 ;

VAN WlLGEN & KRUGF.R 1981 ; Gü.l & GROVES

1984), en Floride (ABRAHAMSON 1984 ; GlVENS

et al. 1984), en Californie (HANES 1970 ;

MALANSON 1984) et concerne aussi bien des

maquis, des garrigues, le chaparral, des landes

(heathlands), des formations herbeuses ou des

formations dominées par des pins. 11 a été quali¬

fié « d’autosuccession » par plusieurs des auteurs

précédemment cités.

Stade initial

Stade intermédiaire

Fig. 9. — Représentation des différentes phases de l'évolution post-incendie des maquis ligno-herbacés.

ADANSONIA, sér. 3 • 1998 • 20(1)

187

Jaffré T., Rigault F. & Dagostini G.

Il apparaît ainsi que le maquis ligne-herbacé des

terrains miniers de Nouvelle-Calédonie est une

formation secondaire adaptée aux incendies. Il

peut être qualifié de « maquis pyrophyte », sa

destruction par le feu étant suivie par un phéno¬

mène de reconstitution, lent en raison des condi¬

tions édaphiques, mais sans changement profond

du type de végétation ni de la flore. 11 s'agit donc

d’un processus de résilience, au sens de HOI.MNG

(1973), qui traduit la capacité du système écolo¬

gique à répondre à une perturbation par le retour

au stade ayant précédé celle-ci.

Le processus étudié correspond au schéma le

plus commun. Il y a lieu cependant de souligner

que, parfois, de nouveaux incendies intervien¬

nent au cours des premières phases de la reconsti¬

tution. C’est notamment le cas lorsque la

présence de Pinus caribaea entraîne localement,

sous son couvert, l’accumulation d’une litière très

inflammable en saison sèche. Celle-ci est formée

d’aiguilles de pins et de frondes desséchées de

Pteridium esmkntum qui prolifère naturellement

sous les pins. L'incendie qui survient alors entraî¬

ne la disparition de certaines composâmes les

plus sensibles de la flore. C’est le cas notamment

de ligneux qui n’ont pas le temps nécessaire pour

accumuler les réserves leur permettant de rejeter

rapidement de souche, et celui des espèces qui

doivent fructifier pour régénérer le potentiel

séminal. Ces disparitions se font toujours au pro¬

fit de Pteridium esculentum qui devient l’espèce

dominante. On est alors en présence d’une fou-

geraie qui apparaît comme le stade ultime de la

dégradation par le feu sur sols ferrallitiques ferri-

tiques moyennement à fortement désaturés sur

substrat d’origine ultramafique. Ce stade, une

fois installé, « s’auro-entretient » en raison de la

forte inflammabilité des frondes sèches produites

chaque année.

Ces différentes phases de l’évolution post¬

incendie des maquis ligno-herbacés sont schéma¬

tisées Figure 9.

Remerciements

Madame Evelyne F A Vif, ainsi que Messieurs Bernard

CERNEAUX et Joseph Favier ont participé aux relevés

de terrain. La dernière phase du travail a bénéficié

d’un crédit CORDET du Ministère des DOM-TOM.

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Manuscrit reçu le 30 décembre 1997;

version révisée acceptée le 13 février 1998.

ADANSONIA, sér. 3 • 1998 • 20(1)

189

Callistemon of New Caledonia transferred

to Mêla leu ca (Myrtaceae)

Lyndley A. CRAVEN

Australian National Herbarium, Centre for Plant Biodiversity Research,

CSIRO Plant Industry, GPO Box 1600, Canberra, ACT 2601, Australia.

craven@pi.csiro.au

John Wyndham DAWSON

School of Biological Sciences, Victoria University of Wellington,

PO Box 600, Wellington, New Zealand.

john.dawson@vuw.ac.nz

KEYWORDS

Myrtaceae,

Callistemon,

Melaleuca,

New Caledonia.

ABSTRACT

The endemic New Caledonian taxa of Callistemon (five species and one

nonautonymic variety) are transferred to Melaleuca. An identification key for

the New Caledonian taxa of Melaleuca is presented.

MOTS CLÉS

Myrtaceae,

Callistemon,

Melaleuca,

Nouvelle-Calédonie.

RÉSUMÉ

Les cinq espèces de Callistemon endémiques de Nouvelle-Calédonie sont

transférées dans le genre Melaleuca. Une clé des Melaleuca néo-calédoniens

est proposée.

ADANSONIA, sér. 3 • 1998 • 20(1)

191

Craven L.A. & Dawson J.W.

Callistemon R. Br. conventionally has been

separated from Melaleuca L. on the basis of the

former genus having the stamens dispersed

around the hypanrhium apex and free staminal

filaments w h créas m Melaleuca the stamens arc in

five groups opposite the petals and ihe filaments

within each group fused proximally for part,

sometimes the greater part, of their length. The

occurrence of 3-groupcd, fused stamens in

Callistemmi has been long known, e.g. Bt.N l HAM

(1867) recorded this condition in the Western

Australian C. glaucus (Bonpl.) Sweet (as G spe-

ciosus DC.) More recentlv, BYRNES (1984) trans-

ferred the eastern Australian C. vinùnalis (Sol. ex

Gaertner) G Don ex Loudon to Melaleuca on

the basis that this species has its stamens in

bundles (i.e. 5-grouped and fused).

Eight species of the Melaleuca-Callistemon com-

plex occur in New Caledonia (DAWSON 1992)

with seven of them endemic and, of the five spe¬

cies attributed to Callistemon, two hâve the sta¬

mens in bundles, i.e. C. buseanus Guillaumin

and G. gnidioides Guillaumin (the latter variably

so, the stamens ranging from free to 5-grouped

and fused). In addition, adjacent stamens occa-

sionally may be fused at rhe filament base in C,

brevisepalus J.W. Dawson and G. suberosus

Panchcr ex Brongn. & Gris. Due to their many

shared features, Dawson (1978) previously had

given a combined gencric description of

Melaleuca and Callistemon in New Caledonia.

In view of the morphological similarities bet-

ween the endemic New Calédonien species of

the complex there is no doubt rhar these belong

to the same genus, They may well be related to

some of the eastern Australian species of

Melaleuca , norably to M. styphcliaidcs Sm. and its

doser relatives; these species in parrictilar hâve

quite similar leaves to several of the New

Caledonian species. The Australian species of

Callistemon are not especially dosely rdated to

the New Caledonian taxa and are presently being

studied by I..A.C. as part of a broader project

dealing with Melaleuca. The New Caledonian

group of species, in our opinion, fies comfortably

within Melaleuca and the necessary transfers of

the six taxa currently ascribcd to Callistemon are

effected below.

The non-endemic New Caledonian species.

Melaleuca ejuiiujuenervia (Cav.) S.T. Bl.ike, occurs

in New Guinea and Australia also (Blake 1968;

CRAVEN in press); it is a memberof the broad-lea-

ved group of paperharks centred upon M. leuca-

dendra (L.) L., rhe type species of Melaleuca.

Melaleuca dawsonii Craven, nom. nov.

Callistemon suberosum Pancher ex Brongn. & Gris,

Bull. .Soc. Bot. France !J: 183 (1864),—Lecrotype:

Pancher s.n„ 1861 (P!), désignaitd by DAWSON, Fl.

Nouvelle-Calédonie et dépendances 18: 230 ( 1992).

Unfortunately, the spécifie epithet “ suberosa ” is

pre-empted in Melaleuca by Al. suberosa

(Schauer) C.A. Gardner and a new epithet is

required in that genus. The new epithet provided

above honours John Wyndham DAWSON, who

has researched New Caledonian Myrtaceae for

many years and co-atithors this contribution.

Melaleuca pancheri (Brongn. & Gris) Craven

& J.W. Dawson, comb. nov.

Callistemon panchcti Brongn. & Gris, Bull. Soc. Bot.

France 11: 183 (1864).—Lecrotype: DepLtnche 513

(Pi), designated by DAWSON, Fl. Nouvelle-

Caledonie et dépendances 18: 234 (1992).

Callistemon suberosum var. mierophyllam Guillaumin,

Mém. Mus. Natl. Flist. Nat. Paris 8: 277 (1962).—

Type: HiirUmann 1389 (holo-, PI).

Melaleuca buseana (Guillaumin) Craven

& J.W. Dawson, comb. nov.

Callistemon busea/mrn Guillaumin, Bull. Mus. Natl.

Flist. Nat. Il: 414 (1939).—Type: Viral s.n. (holo-,

P!).

Callistemon buseanum var. longifolium Guillaumin,

Mém. Mus. Natl. Hist. Nat. Paris 8: 276 (1962),

nom. inval. (Type not designated).

Melaleuca sphaerodendra Craven

& J.W. Dawson, nom. nov.

Callistemon gnidioides Guillaumin, Bull. Soc. Bot.

France 81: 6, 12 (1934).—Type: Cribs 1213 (holo-,

P!).

192

ADANSON1A. sêr. 3 • 1998 • 20(1)

Callistemon of New Caledonia transferred to Melaleuca

A new spécifie epithet is required as “ gnidioides"

is pre-empted in Melaleuca by M. gnidioides

Brongn. & Gris. The epithet selected is derived

from the Greek sphaeret (bail, sphere) and den-

dron (tree) in référencé to the shape of the crown

in many plants of this species. Two varieties are

distinguished:

Melaleuca sphaerodendra var. sphaerodendra

Melaleuca sphaerodendra var. microphylla

(Virot) Craven & J.W. Dawson, comb. nov.

Callistemon gnidioides var. microphyllum Virot, Mém.

Mus. Natl. Hist, Nat., ser. B, Bot. 4: 30 (1953).—

Type: Virot 1313 (holo-, P!).

Melaleuca brevisepala (J.W. Dawson) Craven

& J.W. Dawson, comb. nov.

Callistemon brcvisepalus J.W. Dawson, Fl. Nouvelle-

Calédonie et dépendances 18: 242 (1992).—Type:

MacKee 39544 (holo-, P!).

Key to the New Caledonian species of Melaleuca

3’.

4’.

Leaves 8 mm or more wide.2

Leaves 6 mm wide or less .4

Inflorescences spicate, the groups of flowers or capsules separated by well-marked internodes .

.M. quinquenervia

Inflorescences-capitate, the flowers or capsules densely aggregated and not separated by well-marked inter¬

nodes .....,.. 3

Leafy rwigs 8-11 mm in diameter ..... M. dawsonii

Leafy rwigs 2.5-3.5 mm in diameter ..... M. pancheri

Inflorescences spicate, the groups of flowers or capsules separated by well-marked internodes ...

.........M. buseana

Inflorescences capitale, the flowers or capsules densely aggregated and not separated by well-marked inter¬

nodes . 5

Stamens 25 or more per flower, fused ar the base in 5 groups. (Shrubs in the far south, mostly in stream

beds or swamps) . 6

Stamens 12 or fewer per flower, free or slightly fused at the base in 5 groups. (Shrubs to small trees with

dense roundcd crowns, on well-drained slopes in the north and south) .7

Leaves with évident parallel veins .M. gnidioides

Leaves with obscure parallel veins .M. brongniartii

Stamens longer than the style, capsule glabrous.....8

Stamens shorter than the style, capsule densely pubescent .M. brevisepala

Leaves 3.5-5 mm wide .......M. sphaerodendra var. sphaerodendra

Leaves 2-3 mm wide .M. sphaerodendra var. microphylla

Acknowledgments

The first author thanks Brendan LEPSCHI for

assistance in data collection, Julie MATARCZYK

for literature hunnng and general assistance, and

the Directors and/or Curators of the followtng

herbaria for the opportunity to studv collections

in their care: BRI, CANB, MEL, NSW, P,

WELTU. This research was supported in part by

the Australian Biological Resources Study.

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(Myrtaceae) in northern and eastern Australia, 1.

Austrobaileya 2: 65-76.

ADANSONIA, sér. 3 • 1998 • 20(1)

193

Craven L.A. & Dawson J.W.

Craven L.A. (in press). — Melaleuca, in I. Southwell

(ed.), Behind tbe names: the botany of tea tree, caju-

put and niaouli. Harwood Academie Publishers.

Dawson J.W. 1978.—Pacific capsular Myrtaceae 13.

Melaleuca and Callistemon (New Caledonia).

Blumea 24: 119-122.

Dawson J.W. 1992. — Melaleuca, Callistemon

(Myrtaceae-Leptospermoideae). Flore de la

Nouvelle-Calédonie et dépendances 18: 216-245.

Muséum National d’Histoire Naturelle, Paris.

Manuscript received 28 July 1997;

revised version accepted 9 February 1998.

194

ADANSONIA, sér. 3 • 1998 • 20(1)

Architecture de l’appareil végétatif

et organisation florale

du Dracontium polyphyllum L. (Araceae)

Guylaine POISSON & Denis BARABÉ

Institut de recherche en biologie végétale, Jardin botanique de Montréal,

4101 rue Sherbrooke est,

Montréal, Québec H1X 2B2, Canada.

sema@jonction.net

barabed@ere.umontreal.ca

MOTS CLÉS

Dracontium ,

Araceae,

fleur,

architecture,

inflorescence,

développement.

RÉSUMÉ

Une description de l’appareil végétatif, du développement de la feuille et de

l’anatomie florale du Dracontium polyphyllum est présentée dans le cadre

d’une analyse comparative avec les genres Anchomanes et Amorphnphallus. Du

point de vue architectural, le D. polyphyllum peut être classé dans trois caté¬

gories différentes selon le cadre théorique adopté. Si l'on adopce la classifica¬

tion d’ÉNGLER (1877), cette espèce fait partie du type V de la septième série

qui comprend plusieurs espèces à tubercules. D’autre part, si I on adopte la

classification architecturale d! I AU J: & OLDEMAN (1970), le D. pnlyphyllum

correspond au modèle de Chamberlain. Finalement, si l’on suit la classifica¬

tion de Ray (1987), cette espèce aurait une organisation de type anisophylle

proleptique. La lobation foliaire a un mode de développement marginal et

intramarginal. Dans l'inflorescence, la matutation des fleurs est basipète. La

fleur comprend trois ou quatre carpelles et généralement de 9 à 12 étamines.

KEYWORDS

Dracontium,

Araceae.

flower,

architecture,

inflorescence.

development.

ABSTRACT

A description of the végétative architecture, the development of the leaf and

the floral anatomy of Dracontium polyphyllum is presented in comparison

with the généra Anchonutnes and Amorphophallus. From an architectural

point of view, D. pnlyphyllum can be put in three diffèrent categories depen-

ding on the theoretical Framework chosen. Il we use llic classification of

Engler (1877), this species belongs to the type V of the seventh sériés,

which comprises many tuberous species. On the other hand, if we adopr the

architectural classification of 11 AU P. & OLDEMAN (1970), D. polyphyllum

corresponds to Chamberlain’s model. Finally, if we follow RAY’s classifica¬

tion (1987) this species would bave an organisation of the proleptic aniso-

phyllous type. The foliar lobation has a marginal and intramarginal mode of

development. The flower maturation in the inflorescence is basipetal. The

flower comprises three or four carpels and generaily 9 to 12 stamens.

ADANSONIA, sér. 3 • 1998 • 20 (1)

195

Poisson G. & Barabé D.

INTRODUCTION

Les Araceae comptent plus de 100 genres et

3000 espèces présentant différents types de crois¬

sance, tels que lianescents, épiphytiques aqua¬

tiques ou géophytiques (Hay & MàBBERLEY

1991). Parmi les géophytes, on trouve, entre

autres, les genres Dmcontium avec 23 espèces

néotropicales (Zhu & GRAYUM 1995),

Anchomanes avec 10 espèces africaines et

Amorpbophallus avec 100 espèces paléotropicales

(Grayum 1990).

En général, les espèces de ces genres ne produi¬

sent annuellement qu’une seule inflorescence et

une seule feuille très découpée, pouvant atteindre

jusqu’à 5 m de longueur chez l 'Amotphôphallus

titanum (GANDAW11A1A et al. 1983). ENGLER

(1911) plaçait ces trois genres dans la même

sous-famille, les Lasioideae. Par contre, GRAYUM

(1990) les classe dans trois sous-familles diffé¬

rentes : le genre Dmcontium dans les Lasioideae,

le genre Amorphophallus dans les Aroideae et le

genre Anchomanes dans les Philodendroideae.

Le mode de croissance des géophytes tropicales

en fait un matériel de choix pour l’étude de

l’architecture des plantes herbacées, qui demeure

toujours un domaine peu connu de la morpholo¬

gie, l’architecture de l'appareil végétatif ayant

surtout été analysée chez les arbres (CORNER

1952 j Hallé & OLDEMAN 1970 ; HallË et al.

1978 ; Tomlinson 1962), Rappelons cependant

que les 24 modèles architecturaux décrits pat

Hallé & OLDEMAN ont été repris dans des tra¬

vaux concernant l'architecture des sous-arbustes,

des plantes herbacées et des lianes (Blanc 1978 ;

Cremers 1973, 1974, 1977, 1986, 1992 ;

CREMERS & SELL 1986 ; J LANNODA-ROBINSON

1977 ; N’Diaye 1977 ; Poisson 1996). Engler

(1877) et plus récemment RlTTERBUSCH (1971),

Blanc (1977a,b, 1978, 1980), Ray (1987b,

1988) et Hay (1992) se sont intéressés à l’organi¬

sation de l’appareil végétatif des Araceae sans

approfondir pour autant la question des espèces

géophytiques. Dans cette optique, l’analyse

détaillée de l'organisation végétative et du mode

de croissance du Dracontium polyphyUum amène¬

ra une meilleure compréhension de l'architecture

des plantes géophytiques en général et celle des

Araceae en particulier.

Différents auteurs ont décrit sommairement

l'organisation de l’appareil végétatif du

Dracontium polyphyllum (F.NG1.LR 1877, 1911 ;

Hay Si Mabberlf.y 1991 ; Hay 1992 ; Zhu &

GRAYUM 1995). Néanmoins, ces études se fon¬

dent sur l’analyse d’un ou deux spécimens et ne

tiennent pas compte de la variabilité qui peut

exister entre individus. L’observation de plusieurs

spécimens nous permettra de montra la variabi¬

lité intraspécifique et de faire ressortir les modes

de croissance les plus fréquents.

Cette étude comprend aussi des observations

sur l’anatomie de l’appareil reproducteur.

Plusieurs travaux ont été publiés sur l’organisa¬

tion florale des Araceae (e.g. Eydf. et al. 1967 ;

Hotta 1971 ; Barabé & Forcet 1992 ;

LEHMANN & S Ai i 1ER 1992 ; SCRIBAILO &

Tomlinson 1992 , Barabé & Berirand 1996 ;

Boubes & Barabé 1997). Cependant aucun

d’eux ne traite de façon détaillée de l’anatomie

florale des Dracontium, Afin de combler cette

lacune, il nous a semblé approprié d'inclure

quelques observations sur l'organisation florale

du D. polyphyllum , même si celles-ci ne font pas

l’objet d'une analyse comparative détaillée. Ces

obsetvarions pourront servir dans des études plus

générales portant sur l’ensemble de la famille.

Le présent article a donc pour principal objectif

l’analyse de l’architecture de l'appareil végétatif

du D. polyphyllum en relation avec celle des

genres Amoiphophallus et Anchomanes. Cette ana¬

lyse s’accompagne d’une description des organes

végétatifs et reproducteurs.

MATÉRIEL ET MÉTHODES

Les spécimens de Dracontium polyphyllum utili¬

sés dans cette étude ont été récoltés en Guyane

française, dans la région des montagnes de Kaw,

au sud-est de Cayenne, en mai 1994 et en 1995.

Lors de la récolte, les échantillons ont été fixés au

EA.A. (90 ml d éthanol 70%, 5 ml de formol du

commerce à 35% et 5 ml d’acide acétique).

Microscopie électronique. — Les inflorescences

et les primordiums foliaires ont été observés à

Laide du microscope électronique à balayage

(MEB) JEOL JMS35 du Département de

sciences biologiques de l’Université de Montréal.

196

ADANSON1A. sér. 3 • 1998 • 20(1)

Dracontium polyphyllum L. (Araceae)

Les échantillons choisis ont été préalablement

déshydratés suivant une série ascendante d’étha¬

nol, séchés au point critique avec du CO 9 liqui¬

de dans une bombe Polaron, montés sur supports

métalliques et métallisés h l'or-palladium. Les

observations ont été réalisées à des kilovoltages

différents selon les échantillons (15kV à 25kV),

Microscopie optique. — Pour la microscopie

optique, des inflorescences à différents stades de

développement furent déshydratées suivant une

série ascendante de butanol. Pour faciliter l’orien¬

tation des petits spécimens, nous les avons colo¬

rés à l’éryth rosi ne alcoolique en cours de

déshydratation. Cette étape lut suivie d’un enro¬

bage dans la paraffine selon la technique habi¬

tuelle (GURR 1956). Les coupes sériées (10 pm)

d’inflorescences ont été coloices à la safraninc

1 % (diluée dans de l'eau distillée) et au bleu

d’aniline 1% (dilué dans de l'éthanol 100 %). Les

photomicrographies des coupes furent prises à

l’aide de l'Axiomat Zeiss du Département de

sciences biologiques de l'Université de Montréal.

Diagrammes de l’architecture de l'appareil

végétatif. — Les diagrammes de l'architecture de

l'appareil végétatif des spécimens juvéniles ont

été tracés à partir de la dissection de 8 hulbilles

rapportées de la Guyane française en 1994 et

mises en culture dans les serres du Jardin

Botanique de Monrréaf Quant aux spécimens

adultes, les diagrammes de l’architecture de

l’appareil végétatif ont été réalisés à partir de trois

sources, Sur les 51 plants récoltés, 41 furent dis¬

séqués sur le terrain lors de la récolte, 11 en 1994

et 30 en 1995. Des 10 plants rapportés et conser¬

vés en serre au Jardin Botanique de Montréal, 5

furent disséqués et servirent de modèle pour des¬

siner les diagrammes. Finalement, un plant

conservé dans une mini-serre à température,

humidité et luminosité contrôlées, fut observé

quotidiennement de la floraison à l’apparition de

la feuille annuelle.

RÉSULTATS ET DISCUSSION

Description des appareils végétatif et

REPRODUCTEUR

Tige. — Le tubercule, à face supérieure concave

et à face inférieure convexe, est entouré de

Fig. 1. — Dracontium polyphyllum : A, tubercule (F, base du

pétiole ; Rc, racine contractile) ; B, feuille.

ADANSONIA, sér. 3 • 1998 • 20(1)

197

Poisson G. & Barabé D.

racines contractiles sur sa face supérieure

(Fig. IA). Il peut atteindre 30 cm de diamètre et

20 cm de hauteur chez des spécimens ayant une

feuille de près de 2 m. Lors de la sénescence de la

feuille (Fig. IB), on peut distinguer au sommet

du tubercule le bourgeon apical (Fig. 2A). La

propagation de la plante peut aussi s’efFectuer à

l’aide des bulbilles présentes à la face supérieure

du tubercule. Les bulbilles donneront des plantes

à croissance ntonopodiale.

Feuille. — La feuille composée, en forme de

parasol, comprend un limbe fortement divisé,

soutenu par un long pétiole marbré de violet

(Fig. IB). Son limbe est divisé en trois rachis pri¬

maires de dimension égale (Fig. 2B). Généra¬

lement d’une longueur de 1 m, le pétiole peut

parfois atteindre plus de 2 m. La nervation du

limbe paraît réticulée, semblable à celle des

Dicotylédones, comme chez la plupart des

Lasioideae. F.n règle générale, plusieurs cata-

phylles précèdent la feuille unique produite

annuellement, cependant, chez environ 10% des

spécimens observés, on note la présence de deux

feuilles. Chez les individus portant deux feuilles,

la deuxième occupe la position de la première

cataphvlle et appartient donc au même article

(Fig. 8 F).

Inflorescence. — Le spadice de 4-7 cm de lon¬

gueur et de 1-3 cm de largeur est entouré par

une spathe de 7-15 cm de longueur et de 2-8 cm

de largeur, dont la surface interne est mauve-vio¬

let (Fig. 3A). La spathe, qui se recourbe selon un

angle de 45" à l’apex, ne couvre pas entièrement

le spadice, bien que les marges se chevauchent

(Fig. 3B). L’inflorescence est portée par un

pédoncule de 2 à 15 cm de longueur (ces

mesures incluent la portion souterraine) et de 0,5

à 2 cm de largeur. Nous avons dénombré de 150

à 290 fleurs par inflorescence. À la base de celle-

ci se trouve une cataphvlle qui est souvent plus

longue que le pédoncule. Comme pour les

Lasieae en général, la maturation des fleurs sur

l’inflorescence se fait de façon basipète (Fig. 4A).

Soulignons que ce type de maturation est

contraire à ce qui a été observé chez d’autres

Araceae (BARABK 1994),

Fleur. — La fleur hypogyne se compose de 5, 6

(90% des spécimens observés) ou 7 tépales, géné¬

ralement de 9-12 étamines, quelquefois 7, dispo-

Fig. 2. — Dracontium polyphyllum : A. bourgeon apical au

sommet du tubercule (E, cataphylle) ; B, limbe de la feuille divi¬

sé en trois rachis primaires.

198

ADANSONIA, sér. 3 • 1998 • 20(1)

Poisson G. & Barabé D.

sées sur deux verrieilies, et d’un gynécée trilocu-

laire (90% des spécimens observés) ou tétralocu-

laire (Fig. 4B-C). Chaque loge renferme un ovule

inséré en position axile, dans la partie médiane

de l’ovaire. Il ne semble pas y avoir de règle

concernant la position des gynécées tétralocu-

laires sur l’inflorescence ; ils sont dispersés au

hasard parmi les fleurs à gynécées triloculaires. La

déhiscence des étamines, confinée à la partie

supérieure de l’anthère, est longitudinale. Des

coupes transversales de la partie supérieure de

l’ovaire supère montrent que chaque carpelle

Fig. 4. — Dracontium polyphyllum : A, portion d’inflorescence montrant la maturation basipète des fleurs ; B, fleur à gynécée trilo-

culaire ; C, fleur à gynécée tétraloculaire. (E, étamine ; C, carpelle ; T, tépale).

200

ADANSONIA, sér. 3 • 1998 ■ 20(1)

Dracontium polyphyllum L. (Araceae)

Poisson G. & Barabé D.

possède un canal stylaire qui lui est propre.

Toutefois, dans le style, les canaux se fusionnent

en un seul (Fig, 5A-B). Les pièces florales du D.

polyphyllum se développent dans un sens acropète

(Fig. 4A) : d’abord les tépales, suivis des étamines

et du gynécée. Durant le développement, les

tépales recouvrent progressivement le reste de la

fleur. Contrairement à la plupart des Araceae, le

style du D. polyphyllum est plutôt long, Toutes

les pièces florales renferment des idioblastes à

rapbides d'oxaiate de calcium, comme c’est géné¬

ralement le cas chez les Araceae (Fig. 5C).

Fruit. — La baie de forme obpyramidale mesure

environ 2 cm de longueur et 2 cm de largeur

Dracontium polyphyllum L. (Araceae)

(Fig. 6A) ; le péricarpe plus ou moins dur est vert

foncé. Chaque fruit renferme trois à quatre

graines entourées de mucus.

Graine. — La graine albuminée du D. polyphyl¬

lum est réniforme comme toutes celles des genres

de Lasieae (SeUBERT 1993). Elle arbore un testa

dur de couleur brun-noir, muni de petites

excroissances sur le contour de la partie supérieu¬

re (Fig. 6A. flèche). Elle est convexe sur les deux

côtés et mesure environ 9 mm de largeur, 7 mm

de hauteur et 4 mm d épaisseur.

Pollen. — Le pollen monocolpé de grandeur

moyenne (34 pm), possède une exine réticulée, Il

a une forme de bateau elliptique à symétrie bila¬

térale (Fig. 6B). L’aperture est située au pôle dis¬

tal comme chez les Monocotylédones en général

(Grayum 1992 ; Reille 1990).

Stomates. — Des stomates de type cyclocytique

ont été observés entre les primordiums floraux.

Au sommer de l'inflorescence se trouvent souvent

des groupes de trois ou quatre stomates

(Fig. 6C). Les stomates de la surface inférieure

des feuilles sont paracytiques, tandis que ceux

présents sur le gynécée sont diacytiques selon la

classification de Metcaub &r CHA1.K (1979)

(Fig. 6D).

Croissance et architecture

Tubercule. — Le D. polyphyllum forme générale¬

ment des tubercules orthotropes qui, à la ren¬

contre d’un obstacle, peuvent devenir

pseudo-rhizomateux, plagiotropes. Ceci diffère

de ce qui a été rapporté par KNECHT (1983)

pour le genre Anchomanes. Les espèces de ce

genre produisent un tubercule plagiotrope, où un

nouveau tubercule, issu d'un bourgeon latéral

provenant de Taxe principal apparaît annuelle¬

ment sans que celui de l'année antérieure ne

meure. La croissance du tubercule de D. poly¬

phyllum est monopodiale jusqu'à la maturité

sexuelle ; par la suite, il adopte un mode de crois¬

sance sympodial (Enc.EFR 1877).

Cataphyile. — Comme chez la plupart des

Araceae, l’initiation d'un nouvel axe chez le D,

polyphyllum est associée à la présence de feuilles

spécialisées, les cataphylles ; le limbe est rudi¬

mentaire ou même quelquefois absent. Les pre¬

miers stades du développement des cataphylles

ressemblent à ceux des feuilles, sauf que les pre¬

mières n’atteignent pas 2% de la longueur du

pétiole de la feuille adulte normale. On note une

dégénérescence des tissus du limbe rudimentaire

(Fig, 7A) pendant la période histogène et

d'expansion. Il semble que dans certains cas

(10%), l'une des cataphylles puisse continuer son

développement comme une feuille normale.

Cette deuxième feuille se trouve toujours à la

meme position dans l’organisation de l’appareil

végétatif ; elle suit directement la feuille de

l'année courante (Fig. 8F).

Phase monopodiale, — Les spécimens juvéniles

provenant des bulbilles ont un mode de croissan¬

ce monopodial. Contrairement aux spécimens

adultes, l'apparition de deux feuilles est commu¬

ne chez les plants juvéniles cultivés en serre (45%

des spécimens observés), D après des travaux réa¬

lisées sur les Anthurium et les Philodendron

(Bi ANC 1977a,b), la croissance monopodiale

serait liée au stade juvénile tandis que la croissan¬

ce sympodiale correspondrait au stade adulte

étant donné que chaque article se termine par

une inflorescence Au stade de planrule, la pre¬

mière feuille du D. polyphyllum est pédalée et tri-

partite, comme cela a été rapporté chez les

Amorphophallus. Par contre, chez les Anchomanes,

la première feuille est entière er sagittéc (Knecht

1983). Cependant, selon KNECHT (1983), après

la production d une série de feuilles entières, le

limbe se divise progressivement chez les

Anchomanes pour finalement ressembler à celui

des Dracontium er des Amorphophallus. Chez les

spécimens juvéniles, il est difficile de préciser la

position de la feuille de l’article suivant parmi les

cataphylles, puisque l'ensemble des primordiums

des car.iphylles semblent avoir la possibilité de

devenir une feuille.

Phase sympodiale. — Au cours d'un cycle

annuel, la plante a une période de dormance

entre sa feuillaison et sa floraison. C’est après la

sénescence de la feuille que l’inflorescence appa¬

raît, soit à la fin de la saison des pluies, Chez le

D. polyphyllum , lorsqu’une inflorescence et une

feuille se développent simultanément, cette der¬

nière appartient à l’article suivant, c’est-à-dire

celui qui se développera au prochain cycle phé-

nologique (Tableau 2). Dans ce cas, l’inflorescen¬

ce est à un stade de développement avancé, alors

ADANSONIA, sér. 3 • 1998 • 20(1)

203

Poisson G. & Barabé D.

que la sénescence de la feuille n’a pas encore

débutée. Étant donné que, dans le cycle phénolo-

gique du D. polyphyllum, le développement de la

feuille précède normalement celui de l'inflores¬

cence, cette feuille plus jeune que I inflorescence

ferait donc partie de l’article suivant, Lors de la

dissection du bourgeon terminai, il apparaît clai¬

rement que la structure qui émergera par la suite,

donc celle ayant ie stade de développement le

plus avancé, sera une inflorescence et non une

feuille. Ceci est différent de ce qui à été rapporté

par KnëCHT (1983) pour les Anchomana et les

Amorphophallus où la floraison précède la feuillai¬

son. Inobservation d’un spécimen conservé en

serre appuie notre interprétation. Dans ce cas, le

développement d'une nouvelle inflorescence sui¬

vra la présence simultanée d'une feuille et d’une

inflorescence, et ce, sensiblement à la même date

que pour des plants récoltés dans leur milieu

naturel.

Lobation de la feuille. — Le limbe de la feuille

mature du D. polyphyllum est divisé en trois par¬

ties égales ; celles-ci sont subdivisées en rachis

secondaires, lesquels portent les folioles. Les trois

rachis primaires portent aussi quelques folioles à

leur base, sous le point d’insertion des rachis

secondaires. Sur la jeune feuille encore protégée

par une cataphylle, on observe un rachis central

(antérieur), ainsi que deux rachis opposés l’un à

l’autre (postérieurs), de moindre dimension que

le rachis antérieur (Fig. 7B). Au cours du déve¬

loppement, les deux rachis postérieurs prennent

de l’ampleur pour atteindre des proportions sem¬

blables au rachis antérieur. Ces rachis postérieurs

ne sont pas des rachis secondaires, car chez les

spécimens matures ils sont de forme et de pro¬

portion analogues au rachis antérieur.

Chez le D. polyphyllum , la lobation primaire se

fait marginalement tandis que les dissections

finales des feuilles montrent une lobation d’origi¬

ne intramarginale (Fig. 7B-C). Le processus de

lobation marginal implique l’alternance de

régions de croissance (les folioles) et de régions

sans croissance (les sinus). Par la suite, des sillons

se creusent à une certaine distance de la marge

foliaire pour finalement sc romprent au cours de

l’expansion et ainsi former les folioles (processus

intramarginal). Un mode de développement ana¬

logue se rencontre aussi chez les Palmiers

(Kaplan et al. 1982), À cause de ta présence de

deux processus, le développement de la feuille du

D. polyphyllum diffère de celui d'autres Araceae à

feuille unique très découpée, telles que les

Amorphophallus et les Anchomana , En effet,

d’après Kaplan (1984), chez les Amorphophallus

et la majorité des Araceae à feuilles fortement

disséquées, il n existerait qu’un processus d initia¬

tion marginale. Dans ie cas d' 'Anchomana, Hay

(1992) suggère que seul un processus d initiation

intramarginal interviendrait. Chez, le D, poly¬

phyllum on observe, dans certains cas, des folioles

à fenestration. Cependant l’origine de ces fenes¬

trations semble être une défaillance du processus

intramarginal et non un phénomène de nécrose

comme chez les feuilles à fenestrations typiques

du Monstcra, décrites par Kapi an (1984) et

Melville & Wiuguiy (1969).

La feuille du D. polyphyllum se libère de sa cata¬

phylle protectrice lorsqu'elle est encore à un stade

de croissance peu avancé. Elle complétera donc

son expansion exposée à l’air libre, hors de la

cacaphylle protectrice. Chez le D. polyphyllum,

( expansion foliaire est polaire, c’est-à-dire que le

pétiole débute sa période de croissance sitôt après

l'émergence de la feuille hors de la cataphylle. Ce

processus se fait de façon acropète, tant en lon¬

gueur qu’en diamètre. Durant cette période, le

limbe de la feuille ne subit qu’une légère aug¬

mentation de taille. Les rachis gardent une orien¬

tation similaire à celle qu’ils avaient dans le

bourgeon, c’est-à-dire que le rachis antérieur

pointe vers le sommet et les rachis postérieurs

pointent légèrement vers le bas. Progressivement,

les rachis divergent à angle égal entre eux pour

finalement prendre la forme d’un parasol. Cette

période coïncide avec l'expansion et l’épaississe¬

ment du lirnbe. Cette expansion foliaire tardive

est rapportée pour plusieurs Araceae telles que les

Ancho mânes, Gonaiopus , Anthurium et

Zamiaculcas (Hay & MABBERLF.Y 1991). Par

contre, chez Amorphophallus, dont la feuille est à

première vue semblable à celle du D.

polyphyllum, on note que lors de l’émergence

hors du sol, la feuille est beaucoup plus avancée

dans son processus d’expansion. De plus, chez

VAmorphophallus le processus d'expansion est

plutôt diffus, le pétiole et le limbe de la feuille se

développant de façon synchrone (Hay 1992).

204

ADANSONIA, sér. 3 • 1998 - 20(1)

Dracontium polypbyllum L. (Araceae)

Fig. 7. — Dracontium polyphyllum : A. dégénérescence des tissus du limbe d'une cataphylle ; B, feuille en développement : les

dissections ultimes ont une origine intramarginale ; C, ébauche foliaire ; la lobation primaire est marginale. Ra, rachis central (anté¬

rieur) ; Rp, rachis opposé (postérieur).

ADANSONIA, sér. 3 • 1998 • 20(1)

Poisson G. & Barabé D.

t spadice

spathe

o feuille

o cataphylle

Fig. 8. — Oracontium polyphyllum, diagrammes de l'architecture de l'appareil végétatif : A, spécimen observé quotidiennement de

la floraison à la feuillaison (1995-96) ; B, spécimen disséqué sur le terrain (1994) ; C, 0, spécimens disséqués sur le terrain ( 1994-

95) ; E, spécimen disséqué -sur le terrain (1995) ; F, spécimen à deux teuilles, disséqué sur le terrain (1994-95) , G, spécimen por¬

tant une feuille et une inflorescence simultanément, disséqué sur le terrain (1994-95). Dans l'article de l'année suivante, il n'y a que

deux cataphylles de représentées après la feuille, même si ce nombre peut être variable. — Formule d'embranchement : F,

feuille : C. cataphylle ; Cl .2,3. cataphylle enfouranl l'inllorescence * prochain article, leuîlle. cataphylles et inflorescence de l'article

2 (le chiffre indique le cycle phênologiqua) ; Cb, cataphylle entourant l'inflorescence ; S, spathe ; Sp, spadice, Sf - (cataphylle +

spathe + inflorescence), (le chiffre indique le cycle phénologique). Les parenthèses indiquent une répétition du même organe, le

chiffre suivant la parenthèse indique le nombre de répétitions.

206

ADANSONIA, sér. 3 • 1998 • 20(1)

Dracontium polyphyllum L. (Araceae)

Diagrammes de l’architecture de l’appareil

végétatif des spécimens adultes. —A partir des

47 diagrammes de l’appareil végétatif de spéci¬

mens adultes, nous avons déterminé 5 types

d’organisation (Fig. 8). Le plus fréquent est celui

où l’on dénombre 4 cataphylles sur le même

article après la feuille de l'année courante, c’est-à-

dire celle présente lors de la récolte (Fig. 8C). Ce

mode de croissance sc retrouve chez 54% des

spécimens observés, nonobstant l’année et

l’endroit de récolte Les autres types d’organisa¬

tion observés sont, par ordre d’importance : 5

cataphylles après la feuille chez 28% des spéci¬

mens observés (Fig, 8B), une (9%) (Fig. 8G),

trois (5%) (Fig. 8D), deux (2%) (Fig. 8E) et 6

(2%) (Fig, 8A). Lorsque, sur un meme article, il

n’y a qu'une seule cataphylle après la feuille, on

trouve, chez 50% des spécimens observés, deux

feuilles dans l'année courante (Fig. 8F) et chez

66% une deuxième inflorescence à un stade de

développement avancé (Fig. 8G).

EnGLI-.R (1877) a classé l’organisation de l’appa¬

reil végétatif des Araceae en sept séries. Selon lui,

le D. polyphyllum appartiendrait au type V de la

septième série qui comprend de nombreux

genres à tubercules comme les A morphophallus et

les Anchomanes. La classification de ENGt.FR

repose largement sur la phyllotaxie et sur le type

de croissance (grimpant, rampant, etc,). Dans le

Tableau 1, se trouve détaillée la formule

d’embranchement du D. polyphyllum donnée par

ENGLER. Comme cette formule d’embranche¬

ment est une formule générale qui s'applique

aussi à des espèces munies de plusieurs feuilles, la

concordance exacte avec nos observations ne

peut être absolue. Toutefois, il est possible de

réunir les différentes formules d’embranchements

que nous avons observées (Fig. 8) dans une for¬

mule générale (Tableau 2). Si Ton transpose la

formule du Tableau 2 dans la notation de

ENGLER on obtient une formule qui pourrait à la

rigueur être incluse dans la formule générale de

Engler (Tableau 3).

Le nombre de cataphylles après la feuille de

I article suivant ne peur être déterminé précisé¬

ment lors de la dissection. Le nombre de cata¬

phylles qui se développeront sur l’article du cycle

suivant variera en fonction de facteurs liés à

l'environnement. Par exemple, les spécimens cul¬

tivés en serre ont tous développé un grand

nombre de cataphylles entre la feuille et Tinflo-

resccncc, tandis que ceux disséqués sur le terrain

avaient généralement moins de cataphylles. Ce

phénomène pourrait s'expliquer par le fait que,

lors du transport du tubercule, nous avons pré¬

maturément enlevé la feuille de Tannée courante.

Nous croyons que cette résection aurait provoqué

une augmentation du nombre de cataphylles

dans les articles suivants.

Comme Ta noté Jl ANNODA-Robinson (1977),

certains modèles architecturaux définis chez les

arbres par H ALLÉ &C OLDEMAN (1970) se retrou¬

vent d’une façon miniaturisée chez les herbacées.

Ces auteurs utilisent trois critères architecturaux

pour définir leurs modèles : 1) l orientation de

croissance des axes végétatifs, 2) les caractéris¬

tiques de la croissance des axes, 3) la position des

fleurs ou des inflorescences, Dans le cas du D.

polyphyllum , nous sommes en présence d’une

miniaturisation du modèle de Chamberlain,

comme chez les Araceae en général (Ray 1987b ;

Hay 1992), Dans ce modèle, une succession

indéfinie d’articles, dérivant les uns des aurres par

un mécanisme sympodial, se renouvelle à chaque

cycle de croissance. Chaque article apparaît de

façon acrone à partir d'un bourgeon situé sous

l inflore.scence terminale, Ce modèle ne s'appli¬

querait qu imparfaitement aux Amorphophallus

qui correspondent au modèle mixte de

Tableau 1. — Formule d'embranchement de l'organisation de l'appareil végétatif, 7ème série, type v de la classe

des espèces à tubercules souterrains, selon Engler (1877). N, cataphylle ; L, feuille ; N(n-1), cataphylle qui entoure

l'inflorescence et l’article suivant [next higher shoot order) ; Nn, cataphylle qui entoure l’inflorescence ; S, spathe ;

Sp, spadice. Les parenthèses indiquent une répétition possible de l'organe.

7e série, Type V :

N,N.N,.<l,L),L,N.N.N(n-1),Nn.S,Sp.

N,N,N...(L,L),L,N,N.N(n-1),Nn,S,Sp.

ADANSONIA, sér. 3 • 1998 • 20(1)

207

Poisson G. & Barabé D.

Tableau 2. — Formule d'embranchement de l'organisation de l'appareil végétatif du Dracontium polyphyllum. F =

feuille ; C = cataphylle ; Cl, C2 = cataphylies qui entourent l'Inflorescence et l'article suivant (next higher shoot

order) ; Cb = cataphylle qui entoure l'inflorescence ; S, spalhe ; Sp, spadice. Les parenthèses indiquent une répéti¬

tion du même organe, l'indice donne le nombre de répétitions possible.

(F)1-2-(C) m -C 1 -Cb-S-Sp

Tableau 3. — Formule d’embranchement du Tableau 2 transposée dans la notation de Engler (1877).

N,N,(L),L, N,N...N(n-1),Nn, S.Sp.

_ N,N,L,N,N ,.N(n-1), Nn, S,Sp.

Chamberlain/Tomlinson (BLANC 1978, 1986).

Comme dans le modèle de Chamberlain, les

Amorphophflllm se caractérisent par une succes¬

sion indéfinie d’articles qui se renouvellent à

chaque cycle de croissance et dérivent les uns des

autres par un mécanisme de croissance

sympodial ; cependant, plutôt que de provenir

du sommet de l'article precedent, le nouvel

article émerge de sa base. Cette caractéristique

propre au modèle de Tomlinson explique l’appel¬

lation du modèle. La différence entre ces deux

genres, au niveau de l’architecture de l'appareil

végétatif, pourrait être liée à des variations dans

leur cycle de croissance. Seule une étude détaillée

de l’architecture de l’appareil végétatif des

Amorphophitllus permettra de vérifier cette hypo¬

thèse. Soulignons cependant qu’il est très délicat

d’émettre des généralités pour le genre

Amorphophallus quand on sait qu’il renferme une

centaine d’espèces, dont très peu ont été étudiées

sous ces aspects. Rappelons, â ce sujet, que PATE

& DlXON (1982) décrivent une espèce monocar-

pique, YAmorphophallus glahra, qui est donc-

conforme au modèle fie I loltrum, selon la classi¬

fication de Haï i l' & Oldeman (1970).

Ray (1987a,b ; 1988) a proposé une classifica¬

tion des différents types d’organisation de l’appa¬

reil végétatif des Araceae. Si I on adopte cette

classification, le D. polyphyllum aurait une orga¬

nisation de type anisophylle proleprique ; aniso-

phylle, puisqu'on trouve chez. cette espèce un

nombre variable de feuilles (incluant les cata¬

phylies) et proleptique, car un nouvel article ne

se développe qu’après une période de repos du

bourgeon apical. D’après Ray (1987b), ce type

d’organisation n’a été, jusqu’à maintenant, obser¬

vé que chez les Philodendron sect. Pteromischum ;

toutefois, selon Ray (1988), cette organisation

existerait aussi chez plusieurs espèces de

Monstera. Le D. polyphyllum peut, dans certains

cas, développer une deuxième feuille simultané¬

ment avec l’inflorescence (Fig, 20G). Dans ces

iras exceptionnels, il semble qu’il adopte momen¬

tanément une organisation de l’appareil végétatif

de type anisophylle sylleptique, puisque le nouvel

article provient d’un bourgeon apical qui n’a pas

été soumis à l’effet d’une période de repos.

Selon TOMLINSON (1982), les espèces d'Arâcées

géophytiques seraient néoténiques. La croissance

du D. polyphyllum est caractérisée par une pério¬

de juvénile oit la feuille est peu disséquée et une

période adulte plus longue et permanente, carac¬

térisée par une croissance sympodiale et une

feuille très découpée (Fig. 4). On observe un type

de croissance semblable chez les espèces

d 'Amorphophttllus et à!Anchometnes. Or, ces

espèces géophytiques ne présentent pas de retard

dans le développement somatique, ni d'accéléra¬

tion de la maturation. On ne peut donc parler de

néoténie, comme le suggère TOMLINSON (1982).

Il faudrait plutôt parler de croissance télesco¬

pique où les unités architecturales sont enchaî¬

nées et répétitives (BARABE 1987).

CONCLUSION

Même si la morphologie des Dracontium res¬

semble à celle des Amorphophallus et des

Anchomanes , ces trois genres présentent certaines

différences tant sur le plan de l’organisation végé-

208

ADANSONIA. sér. 3 • 1998 • 20(1)

Dracontium polyphyllum L. (Araceae)

tative que florale. Chez le D polyphyllum , le

mode de lobation de la feuille est marginal et

intramarginal, tandis que, d'après les données

bibliographiques, il serait uniquement marginal

chez Amorphophallus et uniquement intramargi¬

nal chez Anchomanes. On remarque également

que le processus d’expansion foliaire est polaire

chez le D. polyphyllum alors qu'il serait diffus

chez l' Amorphnphallus. Chez le D. polyphyllum ,

l’expansion de la feuille se démule à un stade de

développement plus précoce que chez

l’ Amorphophallus. Enfin, la feuillaison a lieu

avant la floraison chez le D. polyphyllum, tandis

que ce serait le contraire chez les Amorphophallus

et les Anchomanes.

Du point de vue architectural, le D. polyphyl¬

lum peut être classé dans trois catégories diffé¬

rentes selon le cadre théorique adopré. Selon la

classification d’ENULLR (1877), certe espèce cor¬

respond au type V de la septième série. D’autre

part, si I on adopte la classification architecturale

d’HALIl 6c OlDEMAN (1970), le D. polyphyllum

correspond au modèle de Chamberlain, comme

la plupart des Araceae. Finalement, d'après la

classification de Ray (1987), le D , polyphyllum a

une organisation de type anisophylle proleptique,

ce qui a été peu observé chez les Araceae.

Malheureusement, aucune étude comparative

détaillée de l'organogenèse de la feuille des

Araceae géophytiques n'a encore été réalisée. En

conséquence, il serait intéressant d'effectuer pour

les genres Dracontium, Amorphophallus et

Anchomanes une analyse approfondie de la mor¬

phogenèse foliaire en relation avec le mode de

croissance et l’architecture de l'appareil végétatif.

Remerciements

Nous remercions le Professeur L.uc BROUILLET pour

les nombreux conseils qu’il nous a donnés durant la

réalisation de cette émue et pour l’aide apportée lors

de la récolte des échantillons. Le Professeur Joachim

VlETH a eu l'amabilité de lire une première version de

cet article. Nous remercions MM. Georges Cri MEUS

et Jean-Jacques DE GRANVII I r. pour nous avoir per¬

mis de travailler à l'Herbier de Guyane (Cayenne) et

de demeurer durant quelques jours au campement de

l'ORSTOM sur la piste de Ste-Flie. Nous aimerions

aussi souligner l'aide apportée par Mme Audette

Baloup, M. Jacques BALANCER et M. Jean CERDA

dans l’organisation des excursions ou la récolte de

plantes. Enfin nos remerciements s’adressent à Mme

Louise Pelletier et M. Charles BERTRAND pour leurs

nombreux conseils relatifs aux techniques microsco¬

piques. Ce travail a ère réalisé grâce à des subventions

du Fonds FCAR (Fond pour la formation des cher¬

cheurs et l’aide à la recherche, Québec) et du CRSNG

(Conseil de recherches en sciences naturelles et en

génie du Canada).

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210

ADANSONIA. sér 3 ■ 1998 • 20(1)

Encephalartos ituriensis (Zamiaceae):

an emended description

Piet VORSTER

Department of Botany, University of Stellenbosch,

Private Bag XI, 7602 Matieland, South Africa.

pjvor@maties,sun.ac.za

Roy E. GEREAU

Missouri Botanical Garden, P O. Box 299,

St. Louis, MO 63166-0299, U.S.A.

gereau@mobot.org

KEYWORDS

Encephalartos ituriensis,

Zamiaceae,

D.R. Congo

ABSTRACT

The original description of Encephalartos ituriensis (Zamiaceae) is incomplète,

to some extern inaccurate and based in part on immature material. Based on

subsequently collected material and on observations in situ, an emended des¬

cription with illustrations of the fronds and mature female cônes is pro-

vided.

RÉSUMÉ

La description originale d 'Encephalartos ituriensis (Zamiaceae) est incomplète,

parfois inexacte, et en partie fondée sur un matériel immature. Grâce à des

MOTS CLES échantillons récoltés postérieurement et aux observations de terrain, une des-

Encephalartos ituriensis, ,, . , . , r , i»rn

Zamiaceae cription amendée, avec des dessins de frondes et de cônes femelles murs, est

R.D. Congo présentée.

The original description of Encephalartos itu¬

riensis Bamps &: Lisowski (BâMPS & LlSOWSKl

1990) was fairly detailed, but many of the data

presented there were ambiguous or incomplète,

to the extent that when the second author saw

plants in nature, he thought that they could

represent a different species. The size of the

female cône given in the protologue is incorrect

and the information given concerning the surface

seul ptu ring of the exposed faces of the sporo-

phylls is ambiguous. No information was given

concerning the color of male and female cônes,

the prcsence or absence of a petiole, the indu-

mentum of the cônes, the texture of the leaflets,

and the distribution of teeth on the leaflets, ail

characters considered diagnostic by VOKSTER

(1990 and in press). No illustrations of any part

of the mature frond were given; only a line draw-

ADANSONIA, sér. 3 • 1998 • 20(1)

211

Vorster P. & Gereau R.E.

ing was provided of a sccdling plant with two

leaves. Goodu (1989: 229-231) published

detailed color plates of a wholc plant, leaflets,

cônes, and sporophylls, putportedly of £ itu¬

riensis, in the year befbre rhe valid publication of

the binomial. However, the provenance of the

material illustrared cannot be establishcd.

Goodiî’s habit illustration was based on a photo-

graph of a cultivated plant growing on the shore

of Lake Kivu and said to be front the Ituri Foresr,

The cônes that he illustrated are from a different

plant and were received front a collecter in

Zimbabwe without any Jocality data. GOODF.

(1989: 230) provided no scalc of magnification

for the illustrations of the leaflets, and the

dimensions ol 20 X 12 cm giveu for the female

cône (GOOOU 1989: 231) are obviously incorrect

considering the relat ive size of the stent apex and

frond bases attachée!.

The type material as weil the additional spéci¬

mens cited by Bamts 8c Li.SOWSKl (1990) ail

corne from near Nduye in the northeastern

Démocratie Republic of Congo. After considéra¬

tion of the original description, ailowance for

missing and possibly inaccurate informaüon, and

study of Lisowski 41057 (BR) and herbarium

specimens and photographs of Gereau et al. 5413

(Démocratie Republic of Congo, near Nzaro, ca.

30 km northwest ot Nduye; EA, epu [reference

collection at EpuJu], MO, PRE), we conclude

that ail of these specimens are conspecific.

Therefore, it is clear chat the markedly small

female cottes reporied for H. ituriensis by BAMES

LlSOWSKl (1990) were eilher depauperate or

not fully developcd and that the circuinscription

of the species in the protologue is inadéquate.

Accordingly we supply the following emendcd

description:

Encephalartos ituriensis Bamps &t Lisowski

emend. Vorster & Gereau

Plant arborescent, palnt-like in shape, un-

branched but sometimes suckering from base;

stem to 6 m long and 40-50 cm rhick, usually

procumbent with apical portion erect when more

than 2 m long, covered wiih leal base remains in

a regular pattern, the apex glabrous except for

floccose tawny indumentum on cataphylls.

Fronds numerotts, sprcading-ascending in a

hemispheric crown, straight, dark glossy green,

2-3 m long, sessile; base of rachis covered with

short tawny floccose indumentum; leaflets ca. 80

pairs, gradually reduced to a sériés of prickles

toward base of frond; médian leaflets in plane

view oriented at angle of ca. 60° with rachis and

pointing toward frond apex, opposing leaflets set

at angle of slightlÿ less than 180° to each orher

on circumference of rachis, narrowly oblong-

lanceolate and tapering to acute but not pungent

api ces, not or slightlÿ falcace, with (3-)4-6(-9)

teeth rather evcnly distributed along both mar-

gins and ca. 3 teetli crowded near base of distal

margm, in dried specimens finely corrugated on

abaxial surface and margins (including margins

of teeth), revolute, not or only slightlÿ oveflap-

ping, sortiewhat soft-textured, 19-30 X 2-3 cm

(excluding reeth). Male cône [not scen, descript¬

ion from Bami’S &i- Ei.sowski (1990)] nu m ber

unknown, narrowly ellipsoid, borne on a stout

pednncle to 16 cm long and 1.8 cm thick, color

not recorded but presumed similar to female

cône (see below), glabrous, [in dried State?] ca.

26 x 7 cm; exposed faces of male sporophylls

moderately projecring, slightlÿ drooping at

maturity, ca. 2.5 mm wide, the terminal facet

diflerentiated, rhombic, its diameter ca. 60% of

total horizontal diameter ol exposed lace; female

cônes 1-4, ovoid-cylindrical at maturity, borne

on a pedunclc ca. 10 cm long, initially glaucous-

green becoming yellow-green at maturity, glab¬

rous, ca. 55 X 20 cm; exposed faces of female

sporophylls raised, xomewhar pyramidal, ca. 7 X

4.5 cm, the terminal lacet ofjF-center toward base

ol cône, its diameter 25-33% of total horizontal

diameter ol exposed face, the médian facet diffe-

rentiated, with ridges separaring facets clearly dif-

ferentiated; lacets smooth on faces when fresh,

tuberculatc when dry, irargin ol exposed face of

fresh sporophyll tuberculatc. Seed with sarcotcsta

red when fresh, brown when dry, 37-39 X 21-

28 mm with dried sarcotesta intact, 36-38 x 20-

25 mm with sarcotesta removed.—Fig. 1,2.

GliOCSAPHICAL DISTRIBUTION AND HABITAI’

Encephalartos ituriensis is known with certainty

only from the Ituri Forest in the Démocratie

212

ADANSONIA, sér. 3 • 1998 • 20(1)

1 «>SüV«sJm

Pfeîii v;<J

Fiq. 1. —Encephalartos ituriensis. A, mature female cônes in situ; B, mature female cône; C, plant in habitat.—Ali from Gereau et

al. 5413.

ADANSONIA. sér. 3 • 1998 • 20(1)

Encephalartos ituriensis (Zamiaceae)

Republic of Congo at altitudes of 1 100-1200 m,

from five localities forming an arc over a distance

of some 160 km from near Nzaro south-east-

wards to near Nyankunde, with coordinates 1-

2°N-28-30°E. For considérations of conservation

we do not publicize the localities in greater

detail. Wliilc most of the surrounding area sup¬

ports a rnosaic of rnixed moist and single-domi¬

nant moist semi-evergreen Guineo-Congolian

rain forest (WHlTfc 1983). the habitat of

Encephalartos ituriensis lies outside the forest at

somewhat higher élévations on exposed granitic

dômes that support a xerophytic flora with a

number of disjunct and possibly endemic élé¬

ments (LISOWSKI 1992). At one locality E. itu¬

riensis grows among grass in savanna-like

végétation, but this is apparently a degraded

secondary formation that has been altered by

grazing of domestic livestock.

Phenology

Male cônes are known only from April

(Lisowski 42234), and female cônes only from

March (Gereau et al. 5413) and April (Lisowski

42909). The strong seasonaliiy of the annual

rainfall régime in the Ituri Forest (H ART et al.

1996: 547), and additïonally the edaphic dryness

of the extremely well-drairied sites within this

formation on which Encephalartos ituriensis

occurs, make it probable that growrh and seed

production are regulated by précipitation pat¬

terns. The rnonths of March and April corre¬

spond to the end ol the single dry season and the

beginning of the rains.

Diagnostic f ratures and affinities

Geographically Encephalartos ituriensis occurs

closest to E. whitelockii P.JTL Hurter of western

Uganda, but these species are separated by the

Rift Valley, which probably represents a strong

isolating factor. They share glaucotis-green cônes

that at leasr sometimes turn yellow at maturity

(HURTER 1995), lottg-peduncled male cônes,

more or Icss srnooth and glabrous exposed faces

of female sporophylls, and leaflets that overlap

little or not at ail (Vorster &c Heibloem 1995,

as E. successibus Vorster). Encephalartos white¬

lockii diffère from E. ituriensis in its hard- instead

of soft-textured leaflets without a concentration

of teeth near the base of the distal margin of each

leaflet, and more numerous male cônes, up to 8

borne together.

The only other near-equatorial African species

with soft-textured leaflets is Encephalartos lauren-

tianus De Wild,, from the Kwango River valley

bordering the Démocratie Republic of Congo

and Angola. Tins is a larger species, which can be

distinguished by its larger leaflets (up to 50 X

5 cm), long-pcduncled female cônes, and a red-

dish indumentunt on the cônes of both sexes (DE

WlLDEMAN 1903: 10; 1904: 392; Bois 1907;

Gentil 1904a, b; Lebrun 1930).

Discussion

Our cmended circumscription of Encephalartos

ituriensis c-xcludes the illustrations by Goode

(1989 229-231) for the reasons given above. We

extend the diagnostic characters used to recognize

the species, and include a larger range of morpho-

logical variation than originally described by

Bamps & Lisowski (1990). Much of the additio-

nal morphological variation is due to the plants

from Nzaro (cf. Gereau et al. 5413), which are lar¬

ger and more luxuriant than those at Nduye (cf.

Lisowski 4105/). The Nzaro plants hâve longer

médian leaflets with more acute apices and much

larger female cônes, but the morphological diffé¬

rences between the populations appear insuffident

to justifÿ separate taxonomie status.

Acknowl edgemen ts

The second author thanks John and Terese Hart

for their enthusiaslic support and warm hospitality

during his visits to the 1 turi Forest, and Charlotte

Taylor and Gordon McPherson for crirical reading

of the manuscript. The hrsr author’s research was fun-

ded by the Research Fund of the University of

Stellenbosch.

REFERENCES

Bamps P. 6C LISOWSKI S. 199U.—A new species of

Encephalartos (Zamiaceae) from northeastern Zaire.

Ment. New York Bot. Gard. 57: 152-155.

Bois D. 1907.—Les Encephalartos Lemarinelianus et

Lanrrtitiamis. Rev. Hort. 79: 17"’ 179.

DE WlLDEMAN É. 1903. —Études de systématique et de

géographie botaniques sur la Flore du Bas- et du

ADANSONIA, sér. 3 • 1998 • 20(1)

215

Vorster P. & Gereau R.E.

Moyen-Congo. Ann. Mus. Congo , sér. 5, Bot. 1: 1-80.

DE Wildeman É. 1904.—Les Encephalartos congolais:

386-396, in Notices sur des plantes utiles ou intéres¬

santes de la flore du Congo, vol. 1. Veuve Monnom,

Bruxelles.

GENTIL L. 1904a.— Encephalartos from the Congo.

Gard. Chron., ser. 3, 35: 370-371.

GENTIL L. 1904b.—Les cycadées au Congo. Rev.

Hort. Belge Étrangère 30: 7-9.

Goode D. 1989. —Cycads of Africa. Struik

Winchester, Cape Town.

Hart T.B., Dechamps R., Fornier M. & Ataholo

M. 1996.—Changes in forest composition over the

last 4000 years in the Ituri basin, Zaire: 545-563, in

VAN LIER MAESEN L.J.G., VAN DER BURGT X.M. &

van Medenbach De Rooy J.M.(eds.), The

Biodiversity of African Plants , Proceedings ofthe XTVth

AETFATCongress. Kluwer, Dordrecht.

HuRTER P.J.H. 1995.—Two newly described cycads

from Africa. Phytolagia 78: 409-416.

LEBRUN J. 1930.—Note sur F Encephalartos laurentia-

nus De Wild., de la vallée du Kwango. Rev. Zool.

Bot. Africaines 19: 384-390.

Lisowski S. 1992.—Note floristique de Flturi (Haut-

Zaïre). Fragm. Florist. Geobot. 37: 215-220.

VoRSTER P. 1990.— Encephalartos aemulans

(Zamiaceae), a new species from northern Natal. S.

African ]. Bot. 56: 239-243.

Vorster P. (in press).—A review of the arborescent

tropical species of Encephalartos-, in Chen C.J. (ed.),

Proceedings ofthe Fourth International Conférence on

Cycad Biology. Chinese Academy of Sciences, Beijing.

Vorster P. & Heibloem P. 1995. —Encephalartos

successibus (Zamiaceae): a new species from Uganda.

S. African J. Bot. 61: 347-351.

White F. 1983. —The végétation of Africa-, a descriptive

mernoir to accompany the UNESCO/AETFAT/

UNSO végétation map of Africa. UNESCO, Paris.

Manuscript received 4 September 1997;

revised version accepted 24 February 1998.

216

ADANSONIA, sér. 3 • 1998 • 20(1)

Analyse d’ouvrages — Book reviews

L.J. Dorr. Plant Collectars in Madagascar and the

Comoro Islande. A biographical and bibliographi-

cal guide to individuals and groupe who bave col-

lected berbarium material oj algae, bryopbytes ,

fungi, lichens and vascular plants in Madagascar

and tbe Comoro Jslands. Préface de Césaire

Rabenoro. Introduction de Ghillean PRANCE.

The Trustées, Royal Botanic Gardens, Kew, 1

vol., xlvi + 524 p., ISBN 1 900347 18 0

(1997).

En s’engageant dans cette recherche, sans doute

Laurence J. DORR, alors au Missouri Botanical

Garden et actuellement au département de bota¬

nique de la Smithsonian Institution, n’imaginait

pas l’ampleur de la tâche à accomplir. Il en résul¬

te un ouvrage exceptionnel dans la bibliographie

botanique, même si quelques excellentes contri¬

butions antérieures ont tracé la voie, telles celles

de GacnREAIN pour l’ancienne Indochine (avec

portraits, 1943), de I.RTOUZEY pour le

Cameroun (1968), de IlEIT'KR pour l’Ouest

Africain (1971), ou celle de V'AN STEENIS pour

Flora Malesiana (1950), entre autres.

La présente synthèse prend une plus grande

ampleur du lait qu’il s'agit en fait d’une « bio-

bibliographie •» commentée des récolteurs ayant

opéré sur Je territoire considéré, depuis les ori¬

gines (E. DE FlACOURI 1648) jusqu’au milieu

des années 1990. Après quelques pages de don¬

nées techniques, on trouve une très importante

bibliographie générale pp. xvii-xxxiii, un com¬

mentaire (+ carte) sur les espaces protégés de

Madagascar, une liste des herbiers et collections

xylologiques non citées dans l’ed. 8, 1990, de

l’Index Herbariorum, puis un Index d’abrévia¬

tions et symboles utilisés. L’Index des récolteurs

est présenté alphabétiquement, chaque lettre

s’ouvrant sur la photographie d'un paysage, bota¬

nique oit autre, typique de la région. Pour

chaque récolteur, on trouve (si cela est connu, ou

si c’est la cas) : nom et prénom(s), dates et lieux

de naissance et décès, « cursus », activités bota¬

niques (ou générales), à Madagascar ou dans

d’autres régions, éponymie(s) génériques, réfé¬

rences bibliographiques. Concernant la personne,

références de portraits (au trait, peints, photos),

dates des récoltes à Madagascar et aux Comores,

sources de manuscrits, herbiers où sont déposées

des collections. Bien entendu, les nationalités

sont mentionnées et l’un des grands intérêts de

ces biographies est d’avoir recherché le mieux

possible les contributions des Malgaches à la

connaissance de la flore de leur pays au travers

des récoltes qu’ils ont pu faire eux-mêmes, ou

aider à réaliser. Si les voyageurs-naturalistes ont

une place affirmée, on relèvera que nombre de

forestiers, agronomes, administrateurs, etc.

contribuèrent à réunir les collections botaniques

dont nous disposons aujourd'hui et qui sont

l'indispensable outil de travail pour les connais¬

sances systématiques et l'évaluation de la diversi¬

té floristique (et parfois de ses pertes, parallèle¬

ment aux découvertes faites chaque année). Line

marge blanche en bordure de page a permis

l’insertion de nombreux portraits des récolteurs

mentionnés ; mais il est bien évident que dans ce

domaine, beaucoup de visages demeurent incon¬

nus ; on a même démontré récemment que le

portrait de Ph. COMMERSON (p. 93) ne représen¬

tait pas le naturaliste de l’expédition de

Bougainville ! Les choix sont donc parfois fort

difficiles et c’est ce qui explique en partie cer¬

taines lacunes lorsque les recoupements ne per¬

mettaient pas de parvenir à une certitude raison¬

nable. Excellemment présenté, bien lisible malgré

la densité des textes, l’ouvrage de Laurence J.

Dorr procure un véritable plaisir aux lecteurs

intéressés à l’histoire de la botanique, pas seule¬

ment pour Madagascar et les Comores (!). C'est

aussi un hommage rendu à des investigateurs

parfois oubliés ou méconnus qui ont joué un rôle

dans la trame complexe des explorations. Un

exemple en est donné dans les dernières pages,

avec la liste de plus de 500 collaborateurs des

Services forestiers. La livraison inclut un CD.

ADANSONIA. sér. 3 • 1998 ■ 20(1)

217

On ne peut qu’être reconnaissant à l’auteur et

aux autorités de Kew pour l'édition de cet ouvra¬

ge extrêmement documenté qui, même si des

améliorations peuvent enrichir des éditions ulté¬

rieures, pourra faire regretter à beaucoup que

rien de semblable n’existe pour la France !

G.G. AYMONIN

F.G. Brieger, R. Maatsch & K. Senghas (eds.).

“Rudolf Schlecbter. Die orchideen ", 3rd ed. Band

I/B, 31. Lieferung. Blackwell Wissenschafts-

Verlag, Berlin & Parey Verlag, Hamburg, 1905-

1976 pp., figs, 1866-1947. Paper cover, format

18.8x28.5 cm (1995).

Rudolf SCHLECBTER (1875-1925) was a great

German botanisr and traveller connected with

the famous muséum and botanic garden in

Berlin-Dahlem, At first lie was employed as a

gardner, and later as a curator of the herbarium.

Orchids were a real passion of his life, and he

collected tbein during numerous trips to the tro-

pics. One of his greatest Works is the lamous

“Die Orchideen’’ published in Berlin in 1914,

which con tains a synthesis of ail rhe knowledge

of that tirne on orchids. Soon after

SCHLF.CHTHR's death in 1927, the second, revised

and updated édition of this work was published.

In 1970 Brieger, Senghas & Maai.nch initia-

ted the third édition of “Die Orchideen”, which

is completely different from the First édition and

includes two volumes. The First is a taxonomie

description and consists of threc parts; the

second, published in 1985, dcscribes orchids as

horticultural plants and problems connected

with their cultivation.

“Die Orchideen” is to this day the only taxon¬

omie treatment containing keys for détermina¬

tion of ail généra, and in manv cases also species,

especially the ones of patticulat importance in

cultivation mentioned in this work. “Die

Orchideen” includes numerous, good quality

black and white photographs showing représent¬

atives of mo.st généra. However, the many ink

illustrations are not of equal quality, especially

those published in the First chapters.

Fascicle 31, discussed here, is in fact the Final

portion of part B of the first volume. It includes

the following taxa: 5 généra of subtribe

Ornithocephalinae and also 6 subtribes

(Pachyphyllinae. Pterostemmatinae, Raycaden-

coinae, Lockhartiinae, Trichocentrinae and

lonopsidinae) ol tribe Oncidieae, gathered in an

informai group “Tribella Bipolliniata”, ail be-

longing to subtamily Vandoideae.

Orchids ofsubfamily Vandoideae, which are

more or less related to Oruidium. are the most

numerous group in the neotropics. They are said

to includc about 1500 species belonging to

almost 100 généra. Their classification lias evok-

ed lively discussion among taxonomists for years.

The most recent aitempt to classify members of

the Oncidium-jWiance done by Dr ES S LE R (1993)

seems to be unacceptable, because the author

treats ail these généra within the large subtribe

Oncidiinae. SENGHAS used a more practical

rather than phvlogenetic approach to this pro-

blem, dividing Oncidieae into two sériés of sub¬

tribes based on the number of pollinia, without

paying attention to such important Features in

systematics of Vandoideae as tor example rostell-

urn, tegula and viscidiutn structure. Therefore,

aocording to Senghas, Oncidieae includc among

others Stanhopeinae, orchids that, due not only

to their gynostemium structure but also the

architecture of their végétative parts, seem to be

more closely related to Oncidieae than, for

example, to Maxtllarieae.

However, StNGHASs classification is also differ¬

ent from the one suggested by me (S/LACHETKO

1995), among others, due ro the way in which

he classifies subtribes Ornithocephalinae and

Pachyphyllinae.

The other taxa in the “Tribella bipolliniata”

group were divided by SENGHAS into 13 sub¬

tribes, of which Raycadencoinae was described as

new. The taxonomie concept of those subtribes is

in many cases the sanie as mine, and the différ¬

ences concern the systematic position of several

généra. It is worth rriendoning that SENGHAS

considered ürchidutypus Kraenzl. as a separate

genus, although it is usually treated as a synonym

of Pailiypf/yllum Kuilth.

Part G of the first volume, which is to be publi¬

shed in near future, will include the other taxa of

218

ADANSONIA. sér. 3 • 1998 • 20(1)

Vandoideae, suppléments, an index and a listing

of the literature that was used in the work.

Altogether the third édition of “Die Orchideen”

will contain about 3200 pages witli thousands of

photographs and hundreds illustrations. It will

lie the most comprehensive treatment that lias

ever been published on orchidology.

Unfortunately, like every work published over a

long period of time, it may not contain ail of the

most recent taxonomie news; ol which there is a

lot in this group. I hope that the editors will

express their views on at least sonie of them in

the last part of their work.

D.L. SZLACHETKO

T. Shimizu (ed.). Flora of Nagano Préfecture.

Shinano-mainichi Press, Nagano, japan, 1 vol.,

1735 p., illustr., ISBN 4-7840-9725-2, format

A4 (1997).

Ce volumineux ouvrage, en japonais essentielle¬

ment, est une oeuvre collective réalisée en une

vingtaine d’années sous la direction du Prof.

SHIMIZU avec la collaboration de plusieurs bota¬

nistes japonais de la Société botanique de la

Préfecture de Nagano.

Une première partie introductive comprend :

une collection de belles photographies en couleur

(p. III-X) représentant des plantes vasculaires des

différentes /.ones de végétation (alpine, subalpi¬

ne, montagnarde, collinéenne et aquatique) ; une

préface de T. ShimOîU (p. Xl-XIl) ; des

remarques sur la façon de lire l'ouvrage (p. XIII-

XV) avec une carte des 12 sous-préfectures ; le

sommaire détaillé jusqu’au niveau de la famille

en japonais et latin (p. XV1-XX).

La fraction principale est subdivisée en 4

parties : partie l (p. 1-50) : conditions écolo¬

giques (géographie, géologie, climat, types de

végétation) ; partie 2 (p. 51-124) : historique de

la Flore (botanistes et périodiques ayant étudié

la flore de la Préfecture de Nagano) ; partie 3

(p. 125-1544) : Flore proprement dite (sélection

d’Al gu es principaiemen t plane toniques ;

Bryophytes limitées au genre Sphagnum dont

toutes les espèces sont décrites, figurées et identi¬

fiables A l’aide de clés dichotomiques ;

Ptéridophytes avec clés de détermination des

familles, genres, espèces, dessins d’illustration et

carres de distribution ; Gymnospermes et

Angiospermes étudiées exhaustivement comme

les Ptéridophytes ci-dessus. Les 51 nouveautés

scientifiques de la flore sont énumérées en latin.

1 utilisation d'un ordinateur et de fiches perfo¬

rées pour ta détermination des familles et des

espèces de cette Flore est expliquée. Enfin, partie

4 (p. 1545-1674) : Phytogéographie ; considéra¬

tions concernant le passé et le présent ; guides

botaniques pour les 12 sous-régions de la prélec¬

ture ; monuments naturels). Des index en japo¬

nais et en latin terminent l'ouvrage.

Sont jointes à ce document trois cartes en grand

format, une topographique, une géologique en

couleurs et une des types de végétation égale¬

ment en couleurs.

L’analyse résumée qui vient d'être exposée révè¬

le une somme de connaissances peu ordinaire

pour une Flore locale. Nul doute que les ama¬

teurs et les scientifiques japonais puissent en tirer

le meilleur profit et y trouver des intormations

qu’une Flore plus générale ne pourrait pas leur

donner, en particulier en ce qui concerne la dis¬

tribution géographique des espèces ici minutieu¬

sement reportée à l’aide d’un quadrillage particu¬

lier. Les botanistes occidentaux, s’ils ne peuvent

pas, en raison de la langue, comprendre et utili¬

ser toutes les informations de cette Flore, pour¬

ront cependant admirer les 46 photographies en

couleur de quelques paysages végétaux et plantes

de la préfecture de Nagano sans toutefois en

comprendre les légendes en japonais er, grâce aux

noms latins, se Faire une idée suffisamment préci¬

se de la composition floristique de cette région.

Ils sauront particulièrement gré au directeur de

cette Flore d'avoir rappelé dans la partie histo¬

rique le rôle pionnier de botanistes occidentaux

qui ont commencé à défricher le terrain vierge de

La connaissance botanique du Japon et d’avoir

inséré leur portrait dans les pages de cette Flore.

En résumé, on retiendra que cette Flore de la

Préfecture de Nagano est un bel ouvrage, très

bien documenté et digne de figurer dans route

bibliothèque spécialisée.

J.E. Vidal

ADANSONIA, sèr. 3 • 1998 • 20(1)

219

J. Florence. Flore de la Polynésie française. Ed.

ORSTOM, vol. 1, 393 p., 50 fig., 4 p. photos,

2 cartes (1997).

C’est en 1982 que le territoire de la Polynésie

française demanda à PORSTOM, aujourd'hui

l’Institut Français de Recherche Scientifique pour

le Développement en Coopération, de réaliser

une flore moderne de ces îles. Cette étude fut

confiée à Jacques FLORENCE qui venait d’être

recruté par l’Institut.

La flore de la Polynésie française de Drakf. DEL

Ca.STII.Lu, datant de 1893 et la Flore of South-

Eastern Polynesia de F.B.H. & E.D.H. Brown

réalisée après 1922, ne répondaient plus aux

besoins de connaissances plus précises sur la flore

et la végétation des îles. Un travail important sur

le terrain devait d’abord être réalisé pour réunir

les observations et le matériel d’étude nécessaires.

J. Florence s’y est consacré pendant une dizaine

d’années avec beaucoup de persévérance et de

courage, la prospection des parties montagneuses

des îles n’étant pas de tout repos. Un herbier de

plus de 10,000 échantillons a été constitué qui,

avec les collections plus anciennes existant dans

divers grands herbiers mondiaux, a formé la base

de l’étude floristique,

La flore totale des Phanérogames de la dition

est estimée à 675 espèces, ce qui paraît peu com¬

paré à d’autres îles, il est vrai plus étendues,

comme la Nouvelle-Calédonie ou les Hawaï. Son

intérêt réside dans le fait quenviron les 3/4 des

espèces sont endémiques.

Nous saluons ici avec beaucoup d’intérêt et de

sympathie, la sortie du T’ 1 volume de cette flore,

réalisé entièrement par J. FLORENCE. L’ouvrage

compte 393 pages. Dans l'introduction, l’auteur

brosse le cadre phytogéographique, donnant des

renseignements sur la flore et la végétation des

différents archipels constituant la Polynésie fran¬

çaise. L’histoire de l’exploration botanique des

îles est aussi détaillée, et une bibliographie

concernant surtout la végétation est donnée,

ainsi que la liste des abréviations qui sont utili¬

sées pour l’ensemble de la flore. L’auteur a choisi

de placer à la fin de l’ouvrage une clé des familles

des Dicotylédones, ainsi que le glossaire des

termes botaniques, et la liste des échantillons étu¬

diés avec leur localisation.

Sept familles sont traitées : Cannabaceae,

Cecropiaceac, Euphorbiaceae, Moraceae,

Piperaceae, Ulmaceae, Urricaceae. Au total 113

espèces endémiques, indigènes et naturalisées et

55 espèces culrivées. Les Cannabaceae et

Cecropiaceae ne comprennent chacune qu’une

espèce introduite. Les Euphorbiaceae constituent

une des familles les plus importantes pour les îles

polynésiennes, avec 13 genres, 59 espèces endé¬

miques, indigènes et naturalisées et 25 espèces

cultivées. Le genre Glochidion présente une spé¬

ciation très poussée avec 22 espèces endémiques.

Les Moraceae, avec 7 genres et 8 espèces indi¬

gènes ou naturalisées mais aucune endémique,

ont un moindre intérêt. Les Piperaceae comptent

3 genres et 17 espèces indigènes ou endémiques,

3 espèces naturalisées et quelques espèces culti¬

vées i 10 espèces, toutes des Peperomia, sont

endémiques. Les Ulmaceae forment une petite

famille de 3 genres représentés chacun par une

espèce ; une espèce est indigène et deux sont

endémiques. Enfin, les Urticaceae groupent 11

genres et 24 espèces endémiques, indigènes ou

naturalisés ; deux genres, dont un décrit par

Fauteur, sont endémiques. Des espèces endé¬

miques existent dans les genres Elatostemma,

Pilea et Pipturus .

Les noms des auteurs de taxons sont donnés in

extenso, avec les initiales des prénoms. Il est plus

courant de les abréger, mais l’auteur a voulu évi¬

ter toute confusion, l’inconvénient est d’allonger

singulièrement certaines citations.

En ce qui concerne les espèces, la bibliographie

et les synonymes concernent essentiellement la

région. Les types sont cités chaque fois que pos¬

sible. Les descriptions des espèces sont précises et

complètes sans pour autant être exagérément

longues. Elles sont plus succinctes pour les

espèces seulement cultivées. Sont également

notés : les noms vernaculaires, la répartition,

l’écologie, l’utilisation éventuelle. L’ouvrage est

excellemment illustré par 50 planches au trait,

dues pour la plupart à A. DETLOFF, certaines à J.

Lf.MEUX, et par 2 pages de 19 photos couleurs.

On ne peut que féliciter l’auteur pour le sérieux

de ce travail, ce qui laisse bien augurer de la suite

qui lui sera donnée.

J. Bosser

220

ADANSONIA, sér. 3 • 1998 • 20(1)

J.E. Vidal. Paysages végétaux et plantes de la

Péninsule indochinoise. Agence de la

Francophonie (ACCT), ed. Karthala, 248 p., 1

carte, nombreuses photos couleurs (1997).

La flore et la végétation de la Péninsule indo¬

chinoise sont aujourd’hui assez bien connues,

comme l'écrit J.E. VlDAL dans l introducrion à

son ouvrage. Auparavant, il signale les principales

publications botaniques consacrées à cette région

depuis 1790 (« Flora cochinchinensis ») jusqu’à

nos jours (« Flore du Cambodge, du Laos et du

Vietnam » et « Flora of Thailand »), la plupart

destinées aux spécialistes. Aujourd hui, après de

nombreuses années d incertitude politique, les

pays qui constituent cette péninsule (Thaïlande,

Cambodge, Laos, Vietnam) s’ouvrent de plus en

plus au tourisme et il est heureux que les futurs

visiteurs de cette région aient à leur disposition

un guide en langue Irançaise qui les instruira sur

les végétaux qu’on y rencontre.

La présentation de cet ouvrage est particulière¬

ment réussie. Généreusement illustré de belles

photographies en couleurs (242 au total), sa cou¬

verture est souple, la reliure solide, la rédaction

soignée et la typographie variée. Dans une pre¬

mière partie l’auteur décrit les divers types de

végétation schématisés sur une carre en couleurs

et les principales formations végétales.

Globalement, certaines formations sont ana¬

logues à celles qu'on observe dans d’autres

régions tropicales, en particulier la végétation lit¬

torale et paralittorale (mangrove, forêts maréca¬

geuses, plages sableuses) et la végétation d'eau

douce qui renferment plusieurs espèces pantropi-

cales. Les forêts denses humides de basse altitude

et de montagne sont par contre plus originales

car des espèces particulières à la région

(Dipterocarpus spp. par exemple) y dominent. Les

actions destructrices de l’Homme sur la végéta¬

tion (cultures sur brûlis, exploitation forestière

sans contrôle, utilisation de défoliants en période

de guerre) sont rappelées, ainsi que les actions

protectrices (parcs nationaux, réserves naturelles,

jardins botaniques). Cette première partie est

illustrée de 60 photographies en couleurs de très

bonne qualité qui complètent efficacement les

descriptions données pour ces milieux et permet¬

tent de mieux percevoir la physionomie de ces

paysages végétaux.

Dans une deuxième partie sont décrites et illus¬

trées environ 180 espèces. Un grand nombre

d’entre elles sont soit pantropicales, soit cultivées

dans la plupart des régions chaudes du globe ;

étant les plus communes, ce sont celles que le

visiteur rencontrera le plus souvent et dont il

souhaitera être informé. Cependant, on peut

regretter que l’auteur riait pas présenté un plus

grand nombre de plantes indigènes ; les 3000-

4000 espèces endémiques auraient pu faire

l’objet d’un chapitre particulier faisant mieux res¬

sortir l’originalité de la flore de cette péninsule

estimée à environ 15.000 especes. Ma deuxième

remarque a trait à la citation des noms d auteurs

des taxons : délibérément, et pour simplifier la

présentation, lorsque les noms sont des combi¬

naisons postérieures à la publication du basiony-

mc, J.E. VlDAL na retenu comme auteurs que

ceux qui ont établi ces combinaisons. Cet esca¬

motage de l’auteur du basionyme est contraire à

l’article 49 du Code international de nomencla¬

ture botanique et cette présentation serait à

modifier dans les éditions futures de ce livre.

Une partie importante de I ouvrage concerne les

noms vernaculaires des espèces cirées en cambod¬

gien, laotien, thaïlandais, vietnamien, anglais et

français, avec leurs équivalents scientifiques, ainsi

qu’un index récapitulatif des noms scientifiques

et leurs correspondants vernaculaires.

Les visiteurs des pays qui constituent cette

péninsule et tous les naturalistes qui s’intéressent

à cette région ont désormais à leur disposition un

précieux guide de la végétation, en langue fran¬

çaise. Ecrit pat l'un des meilleurs connaisseurs de

la flore du sud-est asiatique après plus de 50

années de recherches sur les groupements végé¬

taux indochinois (et tout particulièrement ceux

du Laos, sujet de la thèse de doctorat que J.E.

VlDAL a soutenue en 1956), cet ouvrage sera cer¬

tainement très apprécié.

J. Jérémie

ADANSONIA, sér. 3 • 1998 • 20(1)

221

Instructions aux auteurs

Adansonia est une revue internationale de biolo¬

gie végétale, consacrée à l’inventaire, l’analyse et

l’interprétation de la biodiversité des Phanéro¬

games. Elle publie des résultats originaux de

recherches en botanique, particulièrement en systé¬

matique et domaines associés : morphologie, anato¬

mie, biologie, écologie, phylogénie biogéographie...

Les manuscrits, dont le nombre de pages n’est pas

limité a priori, devront suivre rigoureusement les

instructions aux auteurs et seront adressés à la

rédaction :

Adansonia

Laboratoire de Phanérogamie

Muséum national d’Histoire naturelle

16 rue BufFon

F-75005 Paris

Tel. : (33) 01 40 79 33 53

Fax : (33) 01 40 79 33 42

e-mail : jeremief^mnhn.fi

Tout manuscrit non conforme aux instructions

sera retourné pour mise au point. Chaque manus¬

crit sera évalué par au moins deux rapporteurs.

Instructions aux auteurs

Chaque manuscrit soumis (y compris les illustra¬

tions) doit être présenté en trois exemplaires au for¬

mat A4, avec un double interligne et des marges

d’au moins 3 cm ; chaque page sera numérotée. Les

illustrations originales seront jointes au manuscrit

définitif, ainsi qu'une disquette 3.5” de format

Apple Macintoch ou compatible IBM, qui devra

contenir également les tableaux (traitement de texte

Word de préférence). Les auteurs devront respecter

les règles du Code International de Nomenclature

Botanique.

Format et structure

Les manuscrits, écrits en français ou en anglais,

doivent être structurés comme suit :

- titre, si possible bref ; un titre courant doit être

proposé ;

- nom(s) et prénom(s) de(s) auteur(s) suivis de

leur(s) adresse(s) professionnelle(s) et, si possible,

de l'adresse électronique ;

- résumés écrits en français et en anglais, suivis des

mots clés et « key words » ;

- dans le texte courant, n’utiliser les italiques que

pour les taxons de rangs génériques et infra-

génériques ;

- dans le texre courant, les références aux auteurs

seront en majuscules, ex. : DutONÎ (2001),

Dupont (2001, 2002), (Diront 2001 ; Durand

& Dupont 2002), Dupont (2001 : 12), Durand

& Dupont (2002, fig. 2) ;

- dans le texte courant, les références aux illustra¬

tions et aux tableaux de l’article seront présentées

ainsi : (Fig. 1), (Fig. 2A.D). (Fig. 2, 5), (Fig. 3A,

6B). (Fig. 3-6) ; (Tableau 1) ;

- les remerciements seront placés à la fin du texte,

avant les références bibliographiques ;

- les références bibliographiques doivent suivre les

exemples donnés ci-dessous ;

- indiquer dans la marge l'emplacement des illustra¬

tions ;

- lournir les légendes des figures sur une feuille

séparée.

Abréviations

Les périodiques doivent être abrégés selon B-P-H

( Botanico-Periodicttm-Huntianum , 1968) et B-P-H/S

( Botanico-PeriodicuU’i-HuntiitnunilSupplcmentum,

1991).

Les titres des ouvrages doivent c-trc abrégés scion

Taxonomie Li fera tare, ed. 2 (SïAFLEU & Cowan

1976-1988), majs avec les lettres initiales en capi¬

tales.

Les noms d’auteurs de taxons doivent être abrégés

selon Authors of Plant Names (Brum.VIITT &

POWKLL 1992).

Les noms des Herbiers doivenr être abrégés selon

Index Herbariorium, cd. 8 (HoLMGREN,

HOLMCRKN & B ARN in 1990).

Illustrations

Les illustrations doivent être réalisées à l’encre de

Chine ou être lournies en impression laser. Les

photographies, bien contrastées, seront sur fond

noir ou blanc. Elles pourront être regroupées et,

ADANSONIA. sér. 3 • 1998 ■ 20(1)

223

Instructions aux auteurs

dans ce cas, identifiées par une lertre en capitales

(A, B, C, ...). Les planches photographiques, pla¬

cées dans le corps de l’article, doivent être traitées et

numérotées comme des figures. Les illustrations

pourront être assemblées sur une colonne (70 mm)

ou sur route la largeur de la justification (144 mm).

Si possible, les légendes et lettrages ne devraient pas

figurer sur les originaux. Ils seront disposés sur un

calque joint à chaque figure, la rédaction se char¬

geant de les placer. Chaque figure doit comporter

une échelle métrique. Les tableaux et graphiques

doivent pouvoir être imprimés sur une page et res¬

ter lisibles après réduction éventuelle. Des photo¬

graphies en couleur pourront être publiées moyen¬

nant une participation financière des auteurs.

Références bibliographiques

Cronquis 1 A. 1981. — An Integra te d System of

Classification of Flowering Plants. Columbia

Univers! ty Press, New York.

GEESINK R. 1981. — Tephrosieae : 245-260, in

Polhill R.M. & Raven P.H. (eds.), Advances in

Instructions to authors

Adansonia is an international journal of plant

biology, devoted to the invenrory, analysis and

interprétation of Phanerogam biodiversity. It

publishes original results of botanical research, par-

ticularly in systematics and related fields: morpho-

logy, anatomy. biology, ecology, phylogeny, bio-

geography, etc.

Manuscripts, without limitation ol the number of

pages, must conform strictly to the instructions to

authors, and should be sent to the Editor:

Adansonia

Laboratoire de Phanérogatnie

Muséum national d Histoire naturelle

16 rue Bufïon

F-75005 Paris

Tel. : (33) 01 40 79 33 53

Fax : (33) 01 40 79 33 42

e-mail : jcremie@mnhn.fr

Instructions to authors

Manuscripts must be submitted in triplicate in A4

Legume Systematics 1 Royal Botanic Gardens,

Kew.

LtROY J. -F. 1978. — Composition, origin and affi-

nities ol the Madagascar vascular flora. Ann.

Missouri Bot. Gard. 65 : 535-589.

Epreuves et tirés à part

Les épreuves seront adressées à l’auteur ou au pre¬

mier auteur (sauf indication contraire) et devront

être retournées corrigées sous huitaine. Pour chaque

article, 25 tirés à part seront fournis gratuitement

par la rédaction ; des tirés à part supplémentaires

pourront être commandés en utilisant un formulai¬

re joint aux épreuves.

Soumettre un article pour publication dans

Adansonia implique que celui-ci n'ait pas été sou¬

mis dans une autre revue. Les droits de reproduc¬

tion de l’article, y compris les illustrations, sont

réservés à la revue. La reproduction de tout ou par¬

tie de l’article doit faire l’objet d'une demande écri¬

te préalable, adressée à la rédaction.

format, double spaced, with margins of at least

3 cm and ail pages numbered. The original figures

should be senr with the revised manuscript, as well

as a 3.5” Apple Macintoch or IBM-compatible

(Word prefèrently) format, which will also contain

any tables. Papers should follow the International

Code of Botanical Nomenclature.

Format and structure

Papers arc to be written in simple and concise

Frcnçh or Lnglish. They should be organized as

follows:

- a brief tille;

- a suggested running head;

- namos(s) and given name(s) ofauthor(s), followed

by their fuII professional adress(es) and, if possible,

e-mail;

- abstracts (in English and frènch), with key words

and "mots clés";

- rexi with italicized words only for taxa of generic

and sub-generic tanks;

- references to authors in main text should be pre-

224

ADANSONIA, sér. 3 • 1998 • 19(2)

Instructions to authors

sented as follows: SMITH (2001), SMITH (2001,

2002), (Smith 2001; Davis & Smith 2002),

Smith (2001: 12), Davis & Smith (2002, fig. 2);

- référencés to illustrations and tables should be

indicated as lollows: (Fig. 1), (Fig. 2A,D), (Fig. 3A,

6B), (Fig. 3-6); (Table 1);

- keep acknowlcdgcmcnts short and place them at

thc end ot die text, before référencés;

- indicate in the margin the suggested placement of

illustrations;

- give captions to illustrations on a separate sheet.

Abbreviations

Periodicals must be abbreviated according to

B-P-H (. Botanico-Periodicum-Huntianum , 1968)

and B-P-H/S ( Botaniço-Periodicum-Huntianum!

Supplementum, 1991),

Abbreviations of book ritles must follow those in

STAFLEU & Cowan (1976-1988): Taxonomie

Literature, ed. 2, but with the first letters in capitals.

Authors’ nantes must be abbreviated according to

Authors of Plant Naines (Brummitt & Powell

1992).

Abbreviations of herbaria must follow Index

Herbariorium, ed. 8 (Holmgren, HOLMGREN &

BARNETT 1990).

Illustrations

Line drawings must be in lndian ink or high qua-

lity laser printouts. Photographs must be high

contrast and be placed on white or black back-

grounds. These can be groupcd into Figures and

identified by letters A, B, C ... Plates will be consi-

dered as Figures and numbered as such. Arrange

Figures to Fit one (70 mm) or two (144 mm)

columns. Letters, numbers, etc., for each figure, are

to be indicated on an accompanying overlay, not

on the original figure. A scale bar is needed for each

figure. No diagram or table is to exceed one page.

References

CRONQU1ST A. 1981. — An Integrated System of

Classification of Flowering Plants. Columbia

University Press, New York.

Geesink FL 1981.—Tephrosieae: 245-260, in POL-

H1LL R.M. & Raven P.H. (eds.), Advances in

Legume Systematics 1. Royal Botanic Gardons, Kew.

Leroy J.-F. 1978.—Composition, origin and affi-

nities of the Madagascar vascular flora. Ann.

Missouri Bot. Gard. 65: 535-589.

Proofs and ojfprittts

Proofs will be sent to the author (or the First

author) for correction and must be returned within

eight days by priority air mail. Authors will receive

twenty-five offprints free of charge; further offprints

can be ordered on a form supplied with the proofs.

The submission of a manuscript to Adansonia

implies that the paper is not being offered for

publication clsewhere. Copyright of a published

paper, including illustrations, bccomes the property

of the journal. Requests to reproduce material from

Adansonia should be addressed to the editor.

ADANSONIA, sér. 3 • 1998 • 20(1)

225

Mise en page

Noémie de la Selle

Packaging Éditorial

Achevé d’imprimer

sur les presses de l’Imprimerie Durand

28600 Luisant (France)

Dépôt légal n° 10154

Printed on acid-free paper

Imprimé sur papier non acide

Date de distribution: 29 juin 1998

Accredited with the International Association for Plant Taxonomy for the purpose of registration

of new names of vascular plants (excluding fossils)

adansonia

5 ^ Hopkins H.C.F.

A révision of Weinmannia (Cunoniaceae) in Malesia and the Pacific. 1. Introduction and an account <

the species of Western Malesia, the Lesser Sunda Islands and the Moluccas

43 • Hopkins H.C.F.

A révision of Weinmannia (Cunoniaceae) in Malesia and the Pacific. 2. Sulawesi and the Philippines

67 • Hopkins H.C.F.

A révision of Weinmannia (Cunoniaceae) in Malesia and the Pacific. 3. New Cuinea, Solomon Islands,

Vanuatu and Fiji, with notes on the species of Samoa, Rarotonga, New Caledonia and New Zealand

107 • Hopkins H.C.F. & Florence J.

A révision of Weinmannia (Cunoniaceae) in Malesia and the Pacific. 4. The Society, Marquesas and

Austral Islands

131 • B° sser )• & Lobreau-Callen D.

w Landiopsis Capuron ex Bosser, genre nouveau de Rubiaceae de Madagascar

# Valkenburg J.L.C.H., Ketner P. & Wilks C.M.

A floristic inventory and preliminary végétation classification of the mixed semi-evergreen rain forest

in the Minkébé région, North East Gabon

163 — Labat J.-N. & Du Puy D.J.

w Sylvichadsia, a new genus of Leguminosae-Papilionoideae-Millettieae endemic to Madagascar

173 • Rigault F. El Dagostini G.

m Impact des feux de brousse sur les maquis ligno-herbacés des roches ultramafiques de Nouvelle-Calédonie

ioi m Craven L.A. & Dawson j.W.

W __ l Kl __ /— _i_ i _ •.

195 •

Callistemon of New Caledonia transferred to Melaleuca (Myrtaceae)

Poisson G. El Barabé D.

Architecture de l'appareil végétatif et organisation florale du Dracontium polyphyllum L. (Araceae)

Vorster P. & Gereau R.E.

211# Encephalartos ituriensis (Zamiaceae): an emended description

217 • Analyse d'ouvrages — Book reviews

Conception Graphique: Isabel Gautray ISSN : 1280-8571

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