a revision of campnosperma (anacardiaceae) in madagascar
TRANSCRIPT
A revision of Campnosperma (Anacardiaceae) in Madagascar
Armand RANDRIANASOLO & James S. MILLER Missouri Botanical Garden, P.O Box 299, St. Louis, Missouri 63116-0299, U.S.A.
KEYWORDS Campnosperma, A n a c a r d i a c e a e , l e p i d o t e scales,
M a d a g a s c a r .
ABSTRACT The genus Campnosperma Thwaites (Anacardiaceae) is treated as consisting
of four species in Madagascar. Two new species are described, illustrated and
compared with C. micranteium Marchand, the only species described in the
Flore de Madagascar, and the recently described C. parvifolium J.S. Miller &
Randrianasolo. A key and descriptions are provided and distribution, habitat
and phenology are also discussed. Campnosperma lepidotum is distinguished
by its leaf undersurface covered with overlapping lepidote scales, and C.
schatzii is marked by its relatively long pedicel and smaller leaf size.
MOTS CLÉS Campnosperma, A n a c a r d i a c e a e ,
écail les l é p i d o t e s , M a d a g a s c a r .
RÉSUMÉ
Le genre Campnosperma Thwaites (Anacardiaceae) comprend quatre espèces
à Madagascar. Deux nouvelles espèces sont décrites, illustrées et comparées à
C. micranteium Marchand, la seule espèce du genre décrite dans la Flore de
Madagascar, et à C. parvifolium J.S. Miller & Randrianasolo, une espèce
récemment décrite. Une clé de détermination, les descriptions des espèces,
ainsi que des informations sur leurs distribution, habitat et phénologie, sont
données. Campnosperma lepidotum se distingue par la surface inférieure des
feuilles couverte d'écaillés lépidotes superposées, et C. schatzii se différencie
par le pédicelle de la fleur qui est relativement long et ses feuilles qui sont de
petite taille.
ADANSONIA, sér. 3 • 1998 • 20(2) 285
Randrianasolo A. & Miller J.S.
T h e genus Campnosperma comprises 1 0 species
of trees and shrubs , wi th six occurr ing in tropical
Asia a n d one each repor ted from Madagascar, the
Seychel les , P a n a m a , a n d Brazi l . O n l y a s ingle
species o f Campnosperma was r e p o r t e d in t he
F lo re d e M a d a g a s c a r ( P E R R I E R D E L A BATHIE
1 9 4 6 ) , a l t hough a second Malagasy species was
r e c e n t l y d e s c r i b e d f r o m t h e M a r o j e j y m a s s i f
( M I L L E R & R A N D R I A N A S O L O 1 9 9 8 ) . T h e reex
a m i n a t i o n of n u m e r o u s b o t a n i c a l c o l l e c t i o n s
m a d e s ince t h e p r e p a r a t i o n o f P E R R I E R D E L A
B A T H I E ' S ( 1 9 4 6 ) t r e a t m e n t i n d i c a t e s t h a t t h e
Malagasy popula t ions of Campnosperma are best
t reated as four morphologica l ly dis t inct species.
Two of these are newly described. Campnosperma
species are usual ly large c a n o p y trees, a n d can
o c c u p y a d iverse t y p e o f h a b i t a t s , s w a m p s to
forests o n well d r a ined soils, r ang ing f rom sea
level to ca. 1 6 0 0 m ( D I N G H o u 1 9 7 8 ) . T h e
Malagasy species are distr ibuted from the wet and
sandy eastern coastal forests at sea level to pre
m o n t a n e and m o n t a n e rainforests u p to 1 7 0 0 m.
Campnosperma species are t rees, w i t h s imple ,
ent i re , petioled, spiral leaves which on b o t h sur
faces very often have m i n u t e , pel ta te or lobed
scales. T h e y have axillary, panicul i form inflores
cences wi th unisexual or bisexual flowers. T h e i r
mos t dist inctive characters in the family are the
n u m b e r of s tamens which is twice the n u m b e r
of petals wi th the epipetalous ones shor ter than
t hose a l t e r n i p e t a l o u s , a n d the i n c o m p l e t e 2 -
celled ovary which w h e n developed to fruit has
very hard and b o n y endocarp and curved seed
( D I N G H O U 1 9 7 8 ) .
C A M P N O S P E R M A Thwai tes
Hooker's J. Bot. Kew Gard. Misc. 6: 65 (1854).
TYPE.—Campnosperma zeylanicum Thwaites.
Key t o the Malagasy species o f Campnosperma
1. Leaves less than 7 cm long and 3.3 cm wide; shrubs or trees of high elevation (ca. 900 m-1800 m) C. parvifolium
V. Most of the leaves gteater than 7 cm long and 3.3 cm wide; trees of low to mid-elevation (sea level ca. 1200 m) 2
2. Leaves with a dense layer of lepidote scales completely concealing the lower leaf surface or nearly so C. lepidotum
2 ' . Leaves with lepidote scales below, but not forming a dense, continuous layer 3 3. Pedicels glabrous, 0.8-2 mm long; leaves 4.5-14.5 cm long, 1.5-5-5 cm wide C. schatzii 3' . Pedicels pubescent, 0.3-0.5 mm long; leaves 11-25 cm long, 4-9 cm wide C. micranteium
C a m p n o s p e r m a l e p i d o t u m C a p u r o n ex
Randr ianasolo & J .S. Miller, sp . nov.
Haec species a congeneris madagascariensibus
similis sed ab eis lamina foliari subtus sqamis lepi-
dotis imbricatis obtecta differt.
TYPE.—SF 8702, Capuron, M a d a g a s c a r , Antsiranana, forêt orienrale (limite supérieure): Massif de l'Ambohitsitondroina à l'est de Mahalevona ( N W de la presqu'île Masoala), ca. 900 m, 4 Dec. 1953, fl., yg. fr. (holo-, P; iso-, M O , TEF).
Funct ional ly dioecious trees, 1 5 - 3 0 m tall, the
y o u n g branches glabrous. T h e leaves s imple, per
s is tent , co r iaceous , very of ten c lus te red at t he
e n d o f b r a n c h e s ; b lades o b o v a t e , 1 0 - 2 1 . 5 c m
l o n g , 4 . 4 - 1 0 c m w i d e , t h e a p e x s o m e w h a t
r o u n d e d o r e m a r g i n a t e , t h e b a s e d e c u r r e n t
cuneate , the marg in entire, slightly revolute, the
adaxial surface glabrous and very sparsely lepido
te, t he abaxial surface covered comple te ly w i t h
lepidote scales fo rming a dist inct m a r o o n layer,
the venat ion p inna te , b r o c h i d o d r o m o u s , p r o m i
nen t below especially the midvein , the secondary
veins more or less parallel, m o r e perpendicular to
the midvein toward the leaf base; petioles 0 . 8 - 1 4
m m , canal icula te o n the adaxial surface, waxy,
lepidotic and wi th very sparse hairs.
2 8 6 ADANSONIA, sér. 3 • 1998 • 20(2)
Campnosperma (Anacardiaceae)
Inf lorescence a s u b t e r m i n a l p a n i c u l i f o r m raceme, bo rne in the very upper leaf axils, 3 .5-19 c m l o n g , t h e b r a n c h e s d e n s e l y l e p i d o t i c w i t h sparse stellate hairs, each flower sub tended by a d e l t o i d b r a c t , 0 . 5 - 0 . 6 m m l o n g , 0 . 5 - 0 . 6 m m w i d e . F l o w e r s m o r p h o l o g i c a l l y b i s e x u a l b u t apparent ly functionally unisexual, small, pedicels 0 .5-0 .8 m m long; sepals 4 , shallowly deba te , val-vate, 0.7-1 m m long, 1-1.2 m m broad, glabrous or glabrescent o n outs ide; petals 4 , ovate, imbr i cate, 1.2-2 m m long, 1-1.2 m m wide , glabrous; s t amens 8, 4 a l tern ipeta lous a n d 4 epipeta lous ,
0 .7-1.5 m m and 0 .5-0 .8 m m respectively in male flowers, m u c h shor ter and sterile in female flowers, inser ted basally o n the ou te r surface of the disk, f la t tened a n d b r o a d e n e d at t h e base, t he an thers ovate, basifixed, in t rorse , w i th long i tu dinal slits, 0 .2-0 .5 m m long; disk more annu la r and cupil i form in female flower, d ish-shaped and c r e n u l a t e in m a l e f lowers; o v a r y a s y m m e t r i c , densely lepidotic, widely ovate to widely depressed ovate , 0 .5 -0 .8 m m long, 0 .3 -0 .5 m m wide , very reduced a n d sterile in male flowers, the style very shor t or obscure, the s t igma subdiscoidal .
2 mm
5 mm
2 mm
4 cm
Fig. 1 . — C a m p n o s p e r m a lepidotum. A, b ranch with l e a v e s and inf lorescence; B, ma le flower; C, female flower; D, fruits.
ADANSONIA, sér. 3 • 1998 • 20 (2) 287
Randrianasolo A. & Miller J.S.
Y o u n g d r u p e s c o v e r e d w i t h l e p i d o t e scales ,
i n c o m p l e t e l y 2 celled fdled by a single curved
embryo , endocarp very ha rd .—Fig . 1.
E C O L O G Y , P H E N O L O G Y A N D D I S T R I B U T I O N . —
T h i s species occurs in nor theas tern Madagascar,
in rainforest be tween 900 and 1150 m elevation,
where precipi ta t ion can be high as 2 0 0 0 m m per
year. It has been collected in flower in D e c e m b e r
and early January.
T h e genus Campnosperma is charac te r ized by
the p resence o f scales o n the i r surfaces (DlNG
HOU 1978) . T h i s species differs markedly from
the o ther Campnosperma species by the undersur -
face of the leaves be ing densely covered by over
lapping lepidote scales (Fig. 2B) . It is similar to
C. micranteium in leaf size, inflorescence pa t te rn
a n d floral c h a r a c t e r s , however , th i s l a t t e r has
m u c h fewer a n d widely spaced lepidote scales.
P A R A T Y P E S . — M A D A G A S C A R : Cours 3668, Antsiranana: au pied de l'Anjanaharibe, 850 m, 16 Dec. 1950, yg. fr. (P, T A N , T E F ) ; Cours 4777, Itinéraire de Didy à Brickaville, fl. (MO, P); Humbert et al. 24532, 24591, massif de l'Anjanaharibe (pentes et s o m m e t N o r d ) à l ' oues t d ' A n d a p a (hau t e Andramonta, bassin de la Lokoho: Nord-Est), forêt ombrophile sur gneiss et granite, 14°40'S, 49°31'E, 900 m, 10 Dec. 1950-3 Jan. 1951, yg. fr. (MO, P); SF925, Caparon, massif de l'Anjanaharibe (W district Andapa), Campement n° 2, 900 m, 16 Dec. 1950, yg. fr. ( M O , P, T A N , TEF) ; SF 925bis, Caparon, est (confins de centre): massif de l'Anjanaharibe, à l'ouest d 'Andapa, 900 m, 16 Dec. 1950, yg. fr. ( M O , P, TAN, TEF); SF 8719, 8720, Caparon, forêt orientale de c imes : mass i f de Г A m b o h i t s i t o n d r o i n a de Mahalevona ( N W de la presqu' î le Masoala) , vers 1100-1200 m, fl. and fr. (P, TAN, TEF); SF 8856, Caparon, forêt orientale, à sa limite supérieure: massif de Beanjada (au Nord de la presqu'île Masoala), 1150 m, 3 Jan. 1954, fl. (MO, P-2 sheets, TAN, TEF).
C a m p n o s p e r m a micrante ium M a r c h a n d
Des Térébinthacées et de ceux de leurs produits qui sont utilisés en pharmacie: 175 (1869); A. Grandidier, Hist. Nat. Mad., Bot. Atlas II, t. 231 (1886).—Type: Boivin s.n, Madagascar, Toamasina: Sainte Marie, 9 Oct. 1850, fl. (holo-, P!).
F u n c t i o n a l l y d i o e c i o u s t r ee s 1 0 - 2 5 m t a l l ,
y o u n g twigs waxy, glabrous, somet imes wi th len-
ticels. T h e leaves s imple , pers is tent , coriaceous;
blades obovate or oblanceolate , 11-25 c m long,
4 - 9 c m w i d e , t h e a p e x e m a r g i n a t e , r e t u s e or
r ounded , the base decur ren t cuneate , the marg in
enrire a n d revolute, the adaxial surface glabrous,
the abaxial surface wi th evenly dis t r ibured lepi
do te scales, the vena t ion p i n n a t e , b r o c h i d o d r o -
m o u s , t h e m i d v e i n p r o m i n e n t u n d e r s u r f a c e ;
petioles 3-7 m m , shallowly channeled , glabrous.
Inflorescence an axillary or sub te rmina l raceme
p a n i c u l i f o r m , c l u s t e r e d a t t h e v e r y e n d o f
branches (as well as the leaves somet imes) , 9-20
c m l o n g , c o v e r e d d e n s e l y w i t h s t e l l a t e h a i r s
m i x e d w i t h sparse l e p i d o t e scales, each flower
s u b t e n d e d by a de l to id to shallowlly t r iangular
b r a c t , ca . 1 m m l o n g a n d 0 . 5 - 1 m m w i d e .
Flowers morpholog ica l ly bisexual b u t func t ion
ally unisexual, very small ca. 1.5 m m long, pedi
cels 0 . 3 - 0 . 5 ( - l ) m m l o n g , g l a b r o u s ; sepals 4 ,
d e b a t e , t r i angu la r or depressed ova te , valvate ,
glabrous, 0 .5-0.6 m m long and 0.5-1 m m wide;
petals 4 , ovate, imbr ica t e , g l ab rous , 1-1.5 m m
long, 0.5-1 m m wide; s tamens 8, 4 al ternipetal-
ous greater in size, 4 al ternisepalous m u c h shor t
er, 0 .8-1 m m a n d 0.5 m m long respectively in
male flowers, s taminodes ca. 0.8-1 m m long in
f e m a l e f l o w e r s , t h e f i l a m e n t s f l a t t e n e d a n d
b r o a d e r at t he base , t h e a n t h e r s ovate or s u b -
globose, basifixed, introrse, opened wi th longi tu
dinal slits, 0 .3 -0 .5 m m long; disk d i sh-shaped ,
cranulate , ca. 1 m m of d iameter in male flowers,
annu l a r , c u p i l i f o r m , edge c r e n u l a t e , ca. 1-1.5
m m in d i ame te r in female flowers; ovary rud i
m e n t a r y in male bu t ovate, very slightly a symme
tric, 1-1.2 m m long, 0 .8-1 m m broad , covered
wi th lepidote scales in female; the style very shor t
ca. 0 .2 m m long , a l m o s t t e r m i n a l ; t he s t i g m a
subdiscoidal variously lobed.
Very y o u n g drupes covered w i t h lepidote a n d
still bear ing a subdiscoidal stigma, m a t u r e fruits
s l ightly asymét r ie , w ide ly e l l ipsoidal , ca. 1 c m
long a n d ca. 0 .8 c m b r o a d , w i t h less l ep ido te
scales o u t s i d e , i n c o m p l e t e l y d i v i d e d i n t o t w o
cells, filled by a s ingle c u r v e d seed, e n d o c a r p
hard.
E C O L O G Y , P H E N O L O G Y A N D D I S T R I B U T I O N . —
T h i s species occupies similar hab i t a t m u c h like
of tha t C. schatzii b u t m o r e restr icted to lower
elevation (up only to ca. 6 0 0 m) where C. schat
zii ranges u p to ca. 1000 m nearby M o r a m a n g a .
288 AD ANSON I A, sér. 3 • 1998 • 20 (2)
Campnosperma (Anacardiaceae)
Campnosperma micranteium usually flowers from Campnosperma micranteium has leaves large as O c t o b e r to January bu t also recorded wirh flow- rhose of C. lepidotum, bu t differs from the latter ers in April and August , a n d fruits from O c t o b e r by its spaced, smaller-sized lepidote scales on the to February. u n d e r s u r f a c e (Fig. 2 A ) . T h i s species is d i s t i n -
Fig. 2 . — S c a n n i n g Electron mic rographs of lower leaf sur face of Ma lagasy Campnosperma s p e c i e s : A, C. micranteium, s c a t t e r ed lepidote s c a l e s on lower leaf sur face {Humbert 24414, MO), s c a l e bar = 50 urn; B, C. lepidotum, over lapping lepidote s c a l e s on lower leaf sur face (Humbert 24535, MO), s c a l e ba r = 50 urn; C, C. schatzii, individual lepidote s c a l e s (Schatz et al. 3419, MO), s c a l e bar = 5 urn; D, C. parvifolium. Individual lepidote s c a l e s (Miller & Randrianasolo 4423, MO), s c a l e bar = 5 urn.
ADANSONIA, sir. 3 • 1998 • 20 (2) 289
Randrianasolo A. & Miller J.S.
gu i shed from С. schatzii by its relatively sho r t pedicel (0 .3-0 .5 m m (rarely 1 m m ) long versus 0 .8-2 m m long) .
O n e popu la t ion from F o r t - D a u p h i n (SF 3352, Capuron) differs from the typical C. micranteium in its smaller leaf size a n d further s tudy and addi t ional collect ions m a y prove it to represent an addi t ional dis t inct taxon.
M A T E R I A L E X A M I N E D . — M A D A C A S C A R : Humbert 24414, environs de Sambava (Cote Nord-Est), bois l i t t o r aux sur sables, mara is à Raph ia , 1 4 ° 1 5 ' S , 50°09'E, 1-5 m, 28 Nov.-3 Dec. 1950, yg. fr. (MO, P); Humblot 462, 28 Jan. 1885, fl. (MO, P-2 sheets); M. de Lastelle s.п., no locality, fl. (P); Perrier de la Bâthie 3002, Maroantsetra, Bois Côte Est, Aug. 1912, fl. (P); 96 R 233, forêt Andrianrantely, Amboditavolo, canton Lohariandava, district Brickaville, 600 m, 22 Aug. 54, fl. buds ( M O ) ; RN 4528, Rakotoniaina, Ambodiarana, district Toamasina, 11 Dec. 1952, fl. (P-2 sheets, T A N , TEF) ; RN 5341, Rakotoniaina, Ambodiriana, district Toamasina, 12 Feb. 1953, fr. (MO, P); RN 6927, Alfred Martin, Ambodiriana, district Toamasina, 6 Jan. 1955, fl. (P, TAN, TEF); RN 7703, Rakoto Jean de la Croix, Angoedro, district Toamasina, 21 Oct. 1955, fl. (P, TEF); SF 887bis, Capuron, Raphières à Antongompahat ra (au N de S a m b a v a ) , 1 D e c . 1 9 5 0 , yg. fr. (P) ; SF 3352, Mandena, Fort-Dauphin; 5 Apr. 1951, fl. (MO, P-2 shee t s , T A N , T E F ) ; SF 7980, P is te A n o s i b e , M o r a m a n g a , fl. ( M O , P, T A N , T E F ) ; SF 9013, Capuron, forêts o r i e n t a l e s : e n v i r o n s du col d'Antandrokolaka (entre Amboditavolo, bassin de la Fanenahana, et Morafeno, bassin de la Rantabe) , Maroantsetra, 400 m, fr. (P, TAN, TEF); SF 11473, Capuron, Sambirano: Massif du Manongarivo, escarp e m e n t s d o m i n a n t la rive g a u c h e de la basse Antsahankolana, à l'Est d'Analalantsoa, Nov. 1954, fl. (P -2 shee t s , T A N , T E F ) ; SF 14177, Capuron, Tampolo Fénérive-Est, 28 May 1955, fr. (P, TAN, T E F ) ; SF 19810, F i ana ran t soa : A m b o l o m b o r o , Canton Ankarimbelo, districr Fort-Carnot, sur une pente exposition ouest à Vohimampay, 25 Oct. 1960, fr. (P, TAN, TEF).
C a m p n o s p e r m a parvi fo l ium C a p u r o n ex J.S.
Mil ler & Randr ianasolo
Novon 8: 170 (1998).—Type: Miller et al. 3529, Madagascar, Antsiranana: Reserve Naturelle Marojejy, along rhe trail to the summit of Marojejy Est, below the third camp, lichen forest and exposed wind-swept ridges, 1100-1300 m, 14°26'S, 49°15 'E , 10 Oct . 1988, fi. (holo-, M O ; iso-, G, K, NY, P, TAN, TEF, US).
Shrub or small tree (2-) 10-12 m tall, the y o u n g
twigs densely covered wi th a mix ture of lepidote
scales a n d s te l la te ha i r s , la ter g l ab rescen t a n d
s m o o t h or somewha t waxy. T h e leaves persistent,
coriaceous; blades elliptic to obovate , ( 1 . 4 - ) 2 - 6 . 7
c m long, (0.9-) 1 . 5 - 3 . 3 c m wide, the apex retuse
to r o u n d e d , the base ob tu se to cunea te , s o m e
t imes briefly decur ren t a long the pet iole for ca.
1-2 m m , the margin strongly revolute, entire, the
adax ia l su r face g l a b r o u s , waxy, a n d l u s t r o u s ,
somet imes sparsely lepidote , the abaxial surface
glabrous or occasionally wi th sparse stellate hairs,
evenly lepidote (appear ing g land-do t ted) , bu t the
scales n o t over lapp ing , t he v e n a t i o n b r o c h i d o -
d r o m o u s , p r o m i n e n t a n d raised on b o t h surfaces,
t h e s e c o n d a r y ve in s 5 - 9 , pa ra l l e l , o f t en o n l y
slightly m o r e p r o m i n e n t t h a n the dense ret icu
l u m o f t e r t i a r y v e i n s o n r h e l o w e r s u r f a c e ;
petioles 3 - 8 m m long, s tout , canaliculate on the
a d a x i a l s u r f a c e , l e p i d o t e t o d e n s e l y s t e l l a t e -
pubescent .
In f lo rescences b o r n e in t he u p p e r leaf axils,
spa r se ly -b ranched panic les or racemes ( l - ) 3 . 3 -
6 . 3 c m long, the branches evenly to densely lepi
d o t e t o s t e l l a t e , e a c h f lower s u b t e n d e d by a
triangular bract 0 . 5 - 1 m m long. Flowers function
ally unisexual ( the p lan ts d ioec ious) , smal l , o n
pedicels 0 . 5 - 1 . 5 m m long, ca. 1 m m in diameter ;
sepals 4 , i m b r i c a t e , de l t a t e , ca. 0 . 5 m m long ,
lepidote; petals 4 , imbricate , broadly elliptic, 1 .5 -
2 m m long, ca. 1 m m wide , glabrous or wi th an
occasional lepidote scale o n the exterior surface;
s tamens 8 , the filaments inserted at the base of
the disk, flattened a n d b roader at the base, ca.
0 . 5 m m long, the anthers ca. 0.2 m m long; nec
tariferous disk annu la r a n d unevenly lobed; ovary
ovoid, small and partially immersed in the a n n u
lar disk, the st igma subdiscoidal .
F ru i t d r u p a c e o u s , b r o a d l y e l l ipsoid , 7 - 8 m m
long, 5 - 6 m m broad, glabrous.
E C O L O G Y , P H E N O L O G Y A N D D I S T R I B U T I O N . —
T h i s s p e c i e s o c c u r s i n c l o u d f o r e s t s o f
Nor theas te rn Madagascar, at 900 m a n d up . It is
k n o w n only from the Marojejy massif a n d several
nearby m o u n t a i n s of N E Madagascar, where it is
often q u i t e a b u n d a n t o n o p e n , h igh-e leva t ion ,
w i n d - s w e p t r i d g e s . I t f l o w e r s i n O c t o b e r -
N o v e m b e r a n d sets fruits in December .
290 ADANSONIA, sér. 3 • 1998 • 20 (2)
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Campnosperma parvifolium differs f rom t h e
o the r species of the genus in Madagascar in its
m u c h smaller leaves, smaller n u m b e r of second
ary veins, a n d lower leaf surface wi th scattered,
small l ep ido te scales (Fig. 2 D ) . It also has the
smallest flowers of any of the Madagascar species
in the genus.
M A T E R I A L E X A M I N E D . — M A D A G A S C A R : Deroin & Badré 40, Maro je jy R N I 12, rive gauche de la Manan ten ina , versant sud de Beondroka, sylve à lichens, 14°25'S, 49°50'E, 1040 m, 12 Nov. 1989, fl. (MO, P); Humbert et al. 24783, Antsiranana: massif de l'Anjanaharibe (pentes et Sommet Nord) à l'ouest d'Andapa, sylve à lichens, HMO'S, 49°26'E, 1600-1800 m, 10 Dec. 1950 - 3 Jan. 1951, yg. fr. (P); Miller & Randrianasolo 4473, Reserve Naturelle de Maro je jy , wes t e rn s lopes and s u m m i t of M t . Beondroka, lichen forest and open wind-swept ridges, 830-1210 m, 14°27'S, 49°47 'E, 26 Oct . 1989, fl. (MO, TAN); Miller & Randrianasolo 4683, Reserve Naturelle de Marojejy, along the trail to the summit of Marojejy Est, N of Mandena, premontane forest, lichen forest, and exposed wind-swept ridges below the 3rd camp, 900-1300 m, 14°26'S, 49°46'E (MO, P, TAN); Rakotomalaza et al. 871, Reserve Naturelle Intégrale de Marojejy, 10.5 km N W of Manantenina, along tr ibutary at head of Andranomifotra River, Campemen t 4, 1625 m, 14°26 '24"S, 49°44 '30"E ( M O , P, T A N ) ; Rasoavimbahoaka 144 ( M O , P, TAN); SF 936, Capuron, massif de l'Anjanaharibe (W d 'Andapa) , 1600 m, 23 Dec. 1950, yg. fr. (P- 2 sheets, TEF).
C a m p n o s p e r m a schatzii Randr ianasolo & J.S.
Miller, sp . nov.
Haec species a congeneris madagascariensibus pedicello sat longo (0.8-2 mm) et foliis parvis (4.5-14.5 cm longis, 1.5-5.5 cm latis) distinguitur.
T Y P E . — S c h a t z et al. 3619, Madagascar, Toamasina: Station Forestière Tampolo, 9 km N of Fénérive-Est. Plot 2. 17°17'15"S, 49°25'11"E, 27 Nov. 1994, fl. (holo-, M O ; iso-, P, TAN).
Funct ional ly dioecious trees, 5-12 m tall, y o u n g
twigs waxy. T h e leaves s imple, persistent, coria
ceous, blades obovate to elliptic or oblanceolate
to ob long , 4 .5 -14 .5 c m long, 1.5-5-5 c m wide ,
the apex e m a r g i n a t e or re tuse or r o u n d e d , the
base decur ren t cuneate , the marg in entire, slight
ly revolute , the adaxial surface sparsely covered
w i t h l e p i d o t e sca les , o r waxy , g l a b r o u s , t h e
abaxial surface m o r e densely covered wi th lepi
do te scales than the upper surface, also glabrous,
t h e v e n a t i o n p i n n a t e , b r o c h i d o d r o m o u s , p r o
m i n e n t undersurface, the secondary veins more or
less parallel, somet imes m o r e obtuse toward the
leaf base; petioles (0 .3-)0 .5- l c m long, shallowly
canaliculate, wi th lepidote and stellate hairs when
young, glabrescent or glabrous when mature .
Inflorescence an axillary raceme panicul i form at
t h e v e r y e n d o f b r a n c h e s , 4 - 1 4 c m l o n g , t h e
branches very often covered wi th lepidote scales
a n d stellate hairs, somet imes glabrous, each flow
er s u b t e n d e d by a smal l t r i a n g u l a r b r a c t , ca.
1 m m long a n d 0.5 m m b r o a d . Flowers func
tionally unisexual, small, pedicel 0.8-2 m m long,
ca. 0 .4 m m in d iameter , s o m e t i m e s few sparse
l ep ido te scales o n its surface; sepals 4 , d e b a t e ,
valvate, 0.5-1 m m long, 0 .5-1 m m wide, some
t imes sparsely l e p i d o t e o n t he exter ior surface
otherwise glabrous; petals 4 , ovate, imbricate , 1-
1.5 m m long, 0.8-1 m m wide, glabrous; s tamens
8, 4 a l t e r n i p e t a l o u s a n d 4 e p i p e t a l o u s m u c h
shorter, 0.8-1 m m a n d ca. 0.5 m m long respec
tively in male flowers, m u c h shor ter a n d sterile
in female flowers, inser ted at the base of ou te r
surface of the disk, the filaments flattened a n d
b r o a d e r a t t h e b a s e , t h e a n t h e r s s u b g l o b o s e
ovoid, basifixed, introrse, wi th longi tudinal slits,
0 .2 -0 .4 m m long; disk m o r e annu la r a n d c u p -
shaped in female flowers, ca. 0.5 m m high, cra-
nula te , shallowly dish-shaped a n d also cranulate
in m a l e f lowers; ovary depressed ova te , a sym
m e t r i c , ca. 0 . 7 - 0 . 8 m m l o n g a n d ca . 1 m m
broad, densely lepidote, very reduced and sterile
in male flowers, the style very short , the s t igma
four lobed a n d diffuse.
Drupes very widely ovate, 5.5-8 m m long, 5-8
m m broad, covered wi th lepidote scales, i n c o m
pletely 2-celled filled by a single curved e m b r y o
or seed, hard endocarp .—Fig . 3 .
E C O L O G Y , P H E N O L O G Y A N D D I S T R I B U T I O N . —
T h i s species occurs in a w i d e range o f h ab i t a t
from sea level to 1200 m . It is d is t r ibuted from
wet sandy coastal littoral forests to h ighland pre-
m o n t a n e r a in fo res t s . It has b e e n c o l l e c t e d in
flower be tween O c t o b e r a n d December , a n d in
fruit between December to February.
T h i s species is d i s t i n g u i s h e d by its relat ively
ADANSONIA, sér. 3 • 1998 • 20 (2) 291
Randrianasolo A. & Miller J.S.
long pedicel (0 .8-2 m m ) . C o m p a r e d to C. lepi-dotum and C. micranteium its leaf size is m u c h smaller (4 .5-14 .5 cm) , a n d it differs from C. par-vifolium in the length of the decurrency of its leaf base (1-2 m m in C. parvifolium vs. 5-10 m m in
C. schatzii). T h e lower leaf surface of this n e w species seemingly has similar sized lepidote scales as C. micranteium, bu t they are fewer in n u m b e r , and thus more widely spaced (Fig. 2 C ) . C. schatzii is n a m e d in h o n o r of George E. S C H A T Z , whose
Fig. 3 . — C a m p n o s p e r m a schatzii: A, b r anch with l e a v e s a n d inf lorescence; B, in f lorescence; C, f emale flower; D, ma le flower; E, fruits.
292 ADANSONIA, sér. 3 • 1998 • 20 (2)
Campnosperma (Anacardiaceae)
collections and work on the flora of Madagascar
have added greatly to our present knowledge .
PARATYPES.—MADAGASCAR: Louvel 38, Toamasina: Tampina, forêt littorale orientale, 25 Oct. , fi. (P); Louvel 51, forêts côtières de l'Est, fi. (P); McPherson 16545, near Andasibe, forest of Mantadia, beyond graphite mine, on ridge above road from km 14, 18°55'S, 48°25'E, 1200 m, 8 Nov. 1994, fl. (MO, P, T A N ) ; RN 8064, A n t s i r a n a n a : A n t a l a h a , Ambohitralanana, 15 Dec. 1956, fi. (P, TAN, TEF); SF 4269, Fénérive-Est, Tampolo, 27 Nov. 1951, fl. (P , T A N , T E F ) ; SF 6462, T a m p i n a - A m b i l a Lemaitso, 27 Dec. 1952, fr. (P, TAN, TEF); SF8979, Capuron, forêt orientale: bassin de la Fananehana, massif de l 'Androna, vers 600 m, fr. (P-2 sheets, TAN, TEF); SF 9039, Capuron, forêt orientale: environs du col d'Antandrokolaka, (entre Amboditavolo, bassin de la Fananehana et Morafeno, bassin de la Rantabe), vers 550 m, fr. (P, TAN, TEF); SF 11448, Capuron, Centre: massif du Manongarivo: Bekolosy, vers 1200 m, 13 Nov. 1954, (P-2 sheets, TAN, TEF); SF 12478, T a m p o l o , C a n t o n Ampas ina , distr ict Fénérive Est, 3 m, 13 Nov. 1954, fl. (P-2 sheets, T A N , T E F ) ; SF 17816, J a rd in B o t a n i q u e n°21 Tampolo , Fénérive Est, 3 m, 3 Dec. 1957, fl. (P, T A N , T E F ) ; SF 22106, Capuron, Est: forêt de Mangalimaso (sur latérite), à l'ouest de Foulpointe, 23 Nov. 1962, fl. (P, TAN, TEF); SF 26820, Capuron, Ankazomanitra, PK 45 Moramanga Anosibe, Canton Anosibe, district Moramanga, forêt sèche, 9 Nov. 1968, fl. (MO, P, TAN, TEF); SF 28156, Capuron, Est: massif boisé de Marozevo (Beforona), 23 Jan.
1968, fr. (MO, P, TAN, TEF); SF 28463, Capuron, Est (confins du Centre): forêt à mousses et sous-bois herbacé aux environs ouest du village d'Antanandava (P.K. 45 de la route Moramanga-Anosibe) , Nov. 1968, fl. (MO, P, TAN, TEF); Tbouars s.n., no locality, sterile (P).
A c k n o w l e d g e m e n t George S C H A T Z was the first to point out the prob
lems w i t h the t a x o n o m y of Campnosperma in Madagascar and we thank bim for his input. We also thank Simon M A L C O M B E R for his valuable comments, Roy G E R E A U for providing the latin description, Mary M E R E L L O for her help with the S E M work, and to Susan A. M O O R E for the illustrations. We are grateful to the staff of the Laboratoire de P h a n e r o g a m i c Muséum National d 'Histoire Naturelle, Paris, for making specimens available for study, both by loan and during visits.
R E F E R E N C E S
DlNG H O U 1978.—Anacardiaceae. Flora Malesiana, ser. 1, 8: 395-548.
M I L L E R J.S. & R A N D R I A N A S O L O A. 1998.—New taxa and n o m e n c l a t u r a l notes on the flora of the Marojejy massif, Madagascar - II. Anacardiaceae: a new species of Campnosperma. Novon 8: 170-172.
P E R R I E R D E LA B Â T H I E H . 1946.—Anacardiacées. Flore de Madagascar et des Comores 114: 1-85.
Manuscript received 26August 1997; revised version accepted 26June 1998.
ADANSONIA, sér. 3 • 1998 • 20 (2) 293