Endemic families of Madagascar. VII.
A synoptic revision of Leptolaena Thouars sensu
stricto (Sarcolaenaceae)
George E. SCHATZ
Missouri Botanical Garden, P.O. Box 299, St. Louis, MO, 63166-0299, U.S.A.
schatz@mobot.org
Porter P. LOWRY II
Missouri Botanical Garden, P.O. Box 299, St. Louis, MO, 63166-0299, U.S.A.
lowry@mobot.org
Laboratoire de Phanérogamie, Museum national d'Histoire naturelle,
16 rue Buffon, 75005 Paris, France.
lowry@mnhn.fr
Anne-Elizabeth WOLF
Laboratoire de Phanerogamie, Museum national d'Histoire naturelle,
16 rue Buffon, 75005 Paris, France.
aewolf@mnhn.fr
KEY WORDS
Sarcolaenaceae,
Leptolaena,
Madagascar,
conservation.
ABSTRACT
As part of an assessment of the vascular plant families endemic to Madagascar
and the Comoro Islands, a synoptic revision is presented of Leptolaena
Thouars sensu stricto (Sarcolaenaceae). Unresolved questions of generic deli-
mitation among Leptolaena s.s., Mediusella (Cavaco) Hutch., Xerochlamys
Baker, and Sarcolaena Thouars preclude a comprehensive revision of
Leptolaena sensu lato at the present time. Two previously recognized species
are retained (L. multiflora and L. pauciflora), L. cuspidata is resurrected from
synonymy, and five new species are described: L. abrahamii, L. delphinensis,
L. gautieri, L. masoalensis, and L. raymondii. Preliminary assessments of the
conservation status of each species are provided, along with a key to the
species in English and French.
RESUME
Familles endemiques de Madagascar. VII. Une revision synoptique du genre
Leptolaena Thouars sensu stricto (Sarcolaenaceae).
Dans le cadre de revaluation des familles de plantes vasculaires de Madagascar
et des Comores, la revision synoptique du genre Leptolaena Thouars sensu
stricto (Sarcolaenaceae) est presentee. Des questions non resolues concernant
ADANSONIA, sér. 3 • 2001 • 23 (2) : 171-189
© Publications Scientifiques du Museum national d'Histoire naturelle, Paris.
171
Schatz G.E., Lowry II P.P. & Wolf A.-E.
la délimitation générique de Leptolaena s.str., Mediusella (Cavaco) Hutch.,
Xerochlamys Baker, et Sarcolaena Thouars excluent actuellement la réalisation
d'une révision complète de Leptolaena sensu lato. Deux espèces déjà recon-
nues (L. multiflora et L. pauciflora) sont retenues ; Leptolaena cuspidata est
réhabilité, et cinq nouvelles espèces sont décrites : L. abrahamii, L. delphinensis,
L. gautieri, L. masoalensis, et L. raymondii. Une évaluation préliminaire pour
la conservation du statut de chaque espèce est présentée, ainsi qu'une clé à
l'espèce, en anglais et en français.
MOTS CLÉS
Sarcolaenaceae,
Leptolaena,
Madagascar,
conservation.
INTRODUCTION
This is the seventh in our series of synoptic
revisions of genera in Madagascar's endemic plant
families (cf. Lowry et al. 1999, 2000; SCHATZ et
al. 1998, 1999a,b, 2000b), which we are publish-
ing to provide an updated taxonomic framework
for assessing the c. 100 species concerned using
the IUCN Red List threat categories (SCHATZ et
al. 2000a), with the ultimate goal of compiling a
Red Data Book detailing the conservation status
of each species. For the present paper, our origi-
nal intention was to review species circumscrip-
tions within Leptolaena Thouars sensu lato, i.e.,
including both Mediusella (Cavaco) Hutch. and
Xerochlamys Baker, thus adopting the broader
generic concept of CAVACO (1952a,b). However,
reconsideration of the characters that have been
utilized to distinguish the three genera from one
another, as well as to distinguish closely related
Sarcolaena Thouars, has caused us to question the
generic limits among the representative taxa.
Sarcolaena has traditionally been circumscribed
to include those taxa that exhibit prefoliation ver-
nation traces on their leaves, a result of an indu-
plicate-folded condition of the immature leaves
during the bud stage, versus the inrolled condi-
tion in related genera. However, a recently
described species of Sarcolaena entirely lacks ver-
nation traces (S. isaloensis Randrianasolo &
J.S. Mill.), and they are also occasionally absent
in S. eriophora Thouars (RANDRIANASOLO &
Miller 1999). Cavaco (1952a,b) emphasized
the complete dissociation of the pericarp at matu-
rity into silky "malpighian" hairs to distinguish
Sarcolaena from Leptolaena sensu lato. However,
Xerochlamys also exhibits comparable dissociation
of the pericarp at maturity into hairs, and
Leptolaena sensu stricto possesses hairs at the base
and sometimes on the interior walls of the
locules. Stipule morphology would appear to ally
Sarcolaena and Xerochlamys, both of which pos-
sess stipules fused into a single hood-like struc-
ture, versus the free stipules of both Leptolaena
s.s. and Mediusella (Capuron 1970). Stamen
number has also been used to distinguish genera,
with Sarcolaena, Xerochlamys, and Mediusella all
possessing "numerous" stamens, i.e., greater than
20, versus Leptolaena s.s. with purportedly 10 sta-
mens. Closer examination of stamen number in
Leptolaena s.s. for the new taxa described here
reveals variable, possibly species-specific values
ranging from 6-8 in L. masoalensis, 8-10 in
L. abrahamii, 10 in L. delphinensis, 10-12 in
L. gautieri, and nearly consistently 11 in L. ray-
mondii. Recent reexamination of pollen of
Sarcolaenaceae showed that differences among
the four genera were insufficient to support either
the division into subgenera or recognition at the
rank of genus (NlLSSON & RANDRIANASOLO
1999).
In light of the problems involved in assigning
taxa within Leptolaena sensu lato and Sarcolaena
to genera, we have decided to restrict the present
study to Leptolaena sensu stricto, i.e., Leptolaena
subgenus Leptolaena [treated as subgenus
"Euleptolaena" by Cavaco (1951)], which com-
prises a morphologically coherent group that in
all likelihood is monophyletic. Leptolaena s.s. can
be distinguished by its smaller number of sta-
mens, generally smaller flowers, and often
glabrous condition. We have examined the speci-
mens at the major herbaria with important hold-
ings of Malagasy plants (K, MO, P, TAN and
172
ADANSONIA, sér. 3 • 2001 • 23 (2)
Revision of Leptolaena (Sarcolaenaceae)
TEF), and have reviewed the circumscription of
the species in the group as recognized by Cavaco
(1952a,b).
In the treatment of Chlaenaceae (= Sarco-
laenaceae) for the Flore de Madagascar, Cavaco
(1952b; see also Cavaco 1952a) recognized seven
species of Leptolaena, two of which (L. multiflora
and L. pauciflora) belong to the group being
treated here. Cavaco also followed Perrier de la
Bathie (1931) in considering L. cuspidata Baker
to be a variety of L. multiflora, whereas we resur-
rect this taxon to the rank of species.
Recent phylogenetic studies utilizing molecu-
lar sequence data indicate that Sarcolaenaceae
and Dipterocarpaceae are sister taxa within an
expanded Malvales (Alverson et al. 1998;
Bayer et al. 1999). Fossil pollen of Sarco-
laenaceae from the Miocene of South Africa
shows that the family was more widespread in
the past and has become endemic to Madagascar
through extinction elsewhere (Coetzee &
Muller 1984).
As in the previous papers in this series, we
have re-evaluated species circumscriptions by
examining morphological features in combina-
tion with eco-geographic parameters, including
bioclimate (Cornet 1974; Schatz 2000; see
also Lowry et al. 1997, 1998) and geological
substrate (Du Puy & Moat 1996). Our analysis
was facilitated by recent collections made by
Malagasy students conducting field work
for their DEA studies at the Universite
d'Antananarivo as part of the Endemic Plant
Families of Madagascar Project (Schatz et al.
2000a).
Comparative analysis of the available material
of Leptolaena sensu stricto has thus enabled us to
propose the following revised taxonomy, in which
eight species are recognized, five of which are
described as new. For the "Material examined"
cited below under each species, abbreviations are
as follows: FC = Foret Classee, PN = Parc
National, RB = Reserve de la Biosphere, RNI =
Reserve Naturelle Integrale, RS = Reserve
Speciale, and STF = Station Forestiere. A full list-
ing of exsiccatae for each species, with complete
localities and latitude/longitude coordinates, has
been compiled for the Madagascar Conspectus
Project (Schatz et al. 1996), and is available on
the World Wide Web through W3 TROPICOS
(http://mobot.mobot.org/W3T/Search/vast.html).
Images of selected taxa are also available on the Web
at (http://www.mobot.org/MOBOT/Madagasc/
sarcolae.html). Specimen data can also be
accessed through the SONNERAT database at
(http://www.mnhn.fr/base/sonnerat.html).
Geographic coordinates indicated in square
brackets were assigned post facto using available
information on Malagasy place names and
topographic maps, compiled as a gazetteer of
botanical collecting localities in Madagascar
(http://www.mobot.org/MOBOT/research/madagascar/
gazetteer/).
LEPTOLAENA Thouars
Hist. Veg. Isles Austral. Afriq. : 41, pl. 11 (1805).
TYPE. — Leptolaena multiflora Thouars.
Key to the species of the Leptolaena s.s.
1.Leaves small, the largest blades usually < 2 cm long (occasionally to 4.5 cm), broadly ovate to circular or
rhombic; inflorescences composed of 1-5 flowers................................................................7. L. pauciflora
1'.Leaves larger, the largest blade > 3 cm long, lanceolate, obovate or narrowly to broadly ovate or elliptic;
inflorescences composed of 7-50 flowers (2-8 in L. masoalensis) ................................................................ 2
2. Leaf apex obtuse to rounded, sometimes emarginate ................................................................................ 3
2'. Leaf apex acute to acuminate ....................................................................................................................4
3.Involucre urceolate, 3 mm tall, covered with dense granular ferruginous indumentum; sepals exserted
1-1.5 mm beyond involucre; leaves narrowly to broadly elliptic, secondary veins (8-)11-13 per side ............
........................................................................................................................................3. L. delphinensis
3'.Involucre ellipsoid to oblong, 7 mm tall, covered with sparse, short white indumentum; sepals completely
included within involucre; leaves elliptic to slightly obovate, secondary veins usually 8 per side ..................
........................................................................................................................................ 5. L. masoalensis
ADANSONIA, ser. 3 • 2001 • 23 (2)
173
Schatz G.E., Lowry II P.P. & Wolf A.-E.
4. Leaves sparsely to moderately sericeous below, with both short appressed trichomes and longer slightly
raised trichomes often exceeding 1 mm long (older leaves occasionally subglabrous), margins distinctly
revolute ..............................................................................................................................8. L. raymondii
4'.Leaves glabrous below or with indumentum restricted to the midvein, margins weakly revolute to flat,
sometimes minutely thickened..................................................................................................................5
5. Leaf blades with evident indumentum on the midvein below ..................................................................6
5'. Leaf blades completely glabrous (rarely a few trichomes at the base of the midvein below) ........................7
6.Involucre usually < 3 mm tall, densely golden yellow granular farinose, rounded at the base with a distinct
peduncle 1 mm long; petals 7-9 X 1-1.5 mm; leaves lanceolate to ovate, apex distinctly long cuspidate ......
............................................................................................................................................2. L. cuspidata
6'.Involucre 3-3.5 mm tall, densely ferruginous granular farinose, usually also with slightly erect whitish tri-
chomes 0.5 mm long, especially in the lower portion, evenly tapering to a cuneate base, with an indistinct
peduncle < 0.5 mm long; petals 10.5-12 X 1.5-2 mm; leaves ovate to broadly ovate, apex acute to acumi-
nate, rarely cuspidate ..............................................................................................................4. L. gautieri
7.Leaves narrowly ovate, usually strongly falcate and inequilateral, often drying reddish black, apex distinctly
acuminate to cuspidate, largest blades 2.5-4.5(-5.5) cm long; sepals at anthesis equal to involucre teeth or
exceeding them by < 0.5 mm.............................................................................................. 1. L. abrahamii
7'.Leaves elliptic, usually equilateral to somewhat inequilateral, rarely a few somewhat falcate, drying brownish
to olive green, largest blades greater than (3-)5.5 cm long; sepals at anthesis exceeding the involucre teeth by
(0.8-)1-1.8 mm ..................................................................................................................6. L. multiflora
Clé des especes de Leptolaena s.str.
1.Feuilles petites, les plus grands limbes generalement < 2 cm de long (parfois jusqu'a 4,5 cm), largement
ovales a circulaires ou rhombiques ; inflorescence composee de 1 a 5 fleurs ........................ 7. L. pauciflora
1'.Feuilles plus larges, les plus grands limbes > 3 cm de long, lanceoles, obovales ou etroitement ou largement
ovales a elliptiques ; inflorescences composees de 7 a 50 fleurs (2 a 8 pour L. masoalensis) ........................ 2
2. Feuille a apex obtus a arrondi, parfois emargine.......................................................................................... 3
2'. Feuille a apex aigu a acumine .................................................................................................................... 4
3.Involucre urceole, haut de 3 mm, couvert d'un indument dense granuleux et ferrugineux ; sepales depassant
l'involucre de 1-1,5 mm ; feuilles etroitement a largement elliptiques, nervures secondaires (8-)11-13 paires....
........................................................................................................................................ 3. L. delphinensis
3'.Involucre ellipsoidal a oblong, haut de 7 mm, couvert d'un court indument blanc clairseme ; sepales inclus
completement dans l'involucre ; feuilles elliptiques a legerement obovales, nervures secondaires generale-
ment 8 paires ..................................................................................................................5. L. masoalensis
4.Feuilles peu a moderement cireuses dessous, portant a la fois de courts trichomes apprimes, et de plus longs
trichomes legerement dresses, souvent depassant 1 mm de long (feuilles plus agees parfois subglabres),
marges distinctement revolutees ........................................................................................ 8. L. raymondii
4'.Feuilles glabres dessous ou a indument limite a la nervure principale, marges faiblement revolutees a planes,
parfois a peine epaissies ............................................................................................................................ 5
5. Limbe foliaire a indument marque sur la nervure principale a la face inferieure ........................................ 6
5'.Limbe foliaire completement glabre (portant rarement quelques trichomes dessous a la base de la nervure
principale) ................................................................................................................................................ 7
6.Involucre generalement < 3 mm de haut, jaune d'or, densement granulo-farineux, arrondi a la base, avec un
pedoncule distinct de 1 mm de long ; petales 7-9 X 1-1,5 mm ; feuilles lanceolees a ovales, apex longuement
cuspide ................................................................................................................................ 2. L. cuspidata
6'.Involucre de 3-3,5 mm de haut, ferrugineux, densement granulo-farineux, portant generalement des tri-
chomes blanchatres, legerement dresses, plus particulierement dans la partie basse, progressivement attenue
en une base cuneiforme, pedoncule indistinct < 0,5 mm de long ; petales 10,5-12 X 1,5-2 mm ; feuilles
ovales a largement ovales, apex aigu a acumine, rarement cuspide .......................................... 4. L. gautieri
7.Feuilles etroitement ovales, generalement tres dissymetriques et falciformes, devenant souvent noir rougeatre
en sechant, apex distinctement acumine a cuspide, les plus grands limbes longs de 2,5-4,5(-5,5) cm ; sepales
a l'anthese de meme taille que les dents de l'involucre ou les depassant de moins de 0,5 mm ......................
............................................................................................................................................ 1. L. abrahamii
7'.Feuilles elliptiques, generalement symetriques a legerement dissymetriques, rarement falciformes, devenant
brunatres a vert olive en sechant, les plus grands limbes depassant (3-)5,5 cm de long ; sepales a l'anthese
depassant les dents de l'involucre de (0,8-)1-1,8 mm .......................................................... 6. L. multiflora
174
ADANSONIA, ser. 3 • 2001 • 23 (2)
Revision of Leptolaena (Sarcolaenaceae)
1. Leptolaena abrahamii G.E. Schatz & Lowry,
sp. nov.
Haec species a congeneris ad Leptolaenam sensu stricto
pertinentibus foliis glabris anguste ovatis falcatis inaequi-
lateralibus in sicco atrorubentibus, sepalis dentes involucrales
aequantibus veleos < 0.5 mm excedentibus distinguitur.
TYPUS. — Miller, Bradford, Rakotonasolo &
Randrianasolo 8746, Madagascar, Prov. Toamasina,
forest N of Andasibe, along railroad tracks, 18°56'S,
48°25'E, 860 m, 20 Oct. 1996, fl. (holo-, MO!; iso-,
P!, TAN!).
Shrubs to trees (1-)4-15 m tall, trunk to
72 cm dbh, stems short tomentose. Leaves char-
taceous, narrowly ovate, strongly falcate, 2.5-
4.5(-5.5) X 1.3-2.5 cm, reduced in the
inflorescence, glabrous, usually drying reddish
black above, dark brown below, base broadly
acute to ovate, margins minutely thickened and
usually somewhat revolute, apex long acuminate
to subcaudate, venation brochidodromous, dis-
tinct, especially below, secondary veins (12-)14-
22 per side, midrib slightly raised above,
distinctly below; petiole 3-4 mm long, 0.5-
0.8 mm diam., narrowly canaliculate, sparsely
puberulous, stipules caducous, scars evident.
Inflorescence terminal, corymbose, c. 20-
40 flowered, axes with minute stellate indumen-
tum often interspersed with scattered appressed
trichomes to 0.3 mm long; peduncle 1-1.5 mm
long, involucre ellipsoid in bud, urceolate to
narrowly cupulate at anthesis, 3.5-4 mm tall, 3-
3.3 mm in diam., densely ferruginous stellate,
apical margin with 6-7 subequal triangular teeth
0.5-1.2 mm tall, 0.5-1.5 mm broad at the base;
sepals obovate, 4 X 2.5 mm, exserted to a height
equal to the involucre teeth or exceeding them
by no more than 0.5 mm, densely white hirsute
outside, glabrous inside; petals narrowly elliptic,
8-9 X 1.8-2.2 mm, pale yellow, sparsely
sericeous outside, exserted 5-6 mm beyond
involucre teeth at anthesis; stamens 8-10, free to
the base, filament c. 5 mm long, anther horse-
shoe shaped, 0.8 mm tall; ovary ovoid, densely
light brown tomentose, style slender, exserted,
5-6 mm long, stigma hemispherical, crateri-
form, with 3 lobes, c. 2 mm across. Fruit ovoid
to ellipsoid, 4-5 mm tall, 2.5-3.5 mm diam.,
irregularly longitudinally channeled, glabrous,
entirely enclosed within the persistent sepals
and involucre. — Fig. 1.
Leptolaena abrahamii is a shrub to medium-
sized tree that occurs in mid-elevation humid
forest from Analamazaotra-Perinet RS and
around Moramanga NNE to Zahamena RNI,
with populations also recorded farther S at
Ampasinambo and Ranomafana PN (Fig. 2). It
can be recognized by its glabrous, narrowly
ovate, falcate leaves with a distinctly acuminate
to cuspidate apex, which usually dry reddish
black, and its sepals that are equal to the
involucre teeth or exceed them by less than
0.5 mm.
Etymology. — The species epithet honors
Jean Prosper Abraham, one of Madagascar's most
distinguished forestry agents who was stationed
for much of his career at Moramanga and
Perinet, where he collected material of Leptolaena
abrahamii (Service Forestier 21955).
Vernacular names. — Amaninombilahy,
Anjananjana, Fotona, Voandrozana.
Conservation status. — With an extent of
occurrence < 20,000 km2 and five sub-popula-
tions, three of which occur within protected
areas, L. abrahamii is assigned a preliminary sta-
tus of Vulnerable (VU).
PARATYPES. — MADAGASCAR, Prov. Fianarantsoa:
Service Forestier 13718, Sahamaizina, Ampasinambo,
Nosy Varika, [20°31'S, 48°00'E], 18 Jan. 1955, y.fr.,
fr. (P); Service Forestier 14241, Ranovao, Ranomafana
PN, [21°18'S, 47°38'30"E], 13 June 1955, bud, fl.,
y.fr. (P, TEF). Prov. Toamasina: Cours 1206,
Nonokambo, [17°44'S, 48°44'E], 950 m, Nov. 1938,
bud, fl., y.fr. (P); Cours 4230, Bemainty a
Androndramanitra [18°00'S, 48°47'E], 800-850 m,
9 Mar. 1951, fr. (MO, P); Dequaire 27840, Zahamena
RNI, [17°38'30"S, 48°50'E], fl., y.fr. (P); Herb. Inst.
Sci. Madag. 3773, Beravina, Analamazaotra-Perinet
RS, [18°56'S, 48°26'E], 1 June 1951, bud (P, TAN);
Herb. Inst. Sci. Madag. 4230 [= Cours 4230],
Bemainty a Androndramanitra [18°00'S, 48°47'E],
800-850 m, 9 Mar. 1951, fr. (TAN); Herb. Jard. Bot.
Tananarive 2844, Sihanaka, 21 Aug. 1937, bud (P);
Mabberley 813, 3 km SE of Perinet, Analamazaotra-
Perinet RS, 18°55'S, 48°25'E, 850-900 m, 29-30 Mar.
1971, ster. (K, TAN); Mission Schedl37, Betsatsa, pres
Perinet, Analamazaotra-Perinet RS, [18°56'S,
ADANSONIA, ser. 3 • 2001 • 23 (2)
175
Schatz G.E., Lowry II P.P. & Wolf A.-E.
Fig. 1. — Leptolaena abrahamii: A, flowering branch; B, leaf (adaxial surface); C, leaf (abaxial surface); D, flower; E, flower with
petals removed. (A-C, Miller et al. 8746; D-E, Reserves Naturelles 15).
48°26'E], 17 Nov. 1952, ster. (TAN); McPherson
17523, Ambatovy, 18°50'54"S, 48°17'56"E, 1000 m,
1 Mar. 1998, fr. (MO, P, TAN); Morat 3218,
Ambatovy, [18°49'S, 48°18'E], May 1969, bud (MO,
P, TAN); Perrier de la Bathie 5336, Analamazaotra-
Perinet RS, [18°56'S, 48°26'E], 800 m, fr. (K);
Reserves Naturelles 15, Analamazaotra-Perinet RS,
[18°56'S, 48°26'E], 30 Oct. 1936, fl. (MO, P);
176
ADANSONIA, ser. 3 • 2001 • 23 (2)
Revision of Leptolaena (Sarcolaenaceae)
Reserves Naturelles 1587, Sahamalaza, Mana-
kambanihy-Est, Zahamena RNI, [17°38'30"S,
48°50'E], 12 Nov. 1948, bud, fl., y.fr. (P); Reserves
Naturelles 3169, Sahatavy, Zahamena RNI,
[17°38'30"S, 48°50'E], 14 Sep. 1951, bud (MO, P,
TAN); Reserves Naturelles 10953, same locality,
12 June 1960, bud, fl., (P, TEF); Reserves Naturelles
12684, Sahamalaza, Zahamena RNI, [17°38'30"S,
48°50'E], 28 Dec. 1963, fr. (TEF); Schatz &
Rabenantoandro 3991, Analamazaotra-Perinet RS,
18°55'58"S, 48°25'16"E, 16 Mar. 2000, ster. (MO);
Schatz & Rabenantoandro 3993, 0.5 km E of Andasibe,
18°55'11"S, 48°25'17"E, 16 Mar. 2000, fr. (G, K, MO,
P, TEF); Service Forestier 3497, Masse (= Ampasimpotsy-
Gara), Analamazaotra-Perinet RS, [18°58'30"S,
48°21'E], 26 Apr. 1951, bud (P, TAN, TEF); Service
Forestier 3773, Beravina, Analamazaotra-Perinet RS,
1 June 1951, bud, fl. (P, TAN, TEF); Service Forestier
5070, Boqueton Abehovana (Piste Moramanga-
Farizana), [19°05'S, 48°15'E], 14 Apr. 1952, bud (P,
TEF); Service Forestier 6242, Antsahatsaka, Beravina,
Analamazaotra-Perinet RS, [18°58'S, 48°16'30"E],
5 Nov. 1952, fl. (MO, P, TEF); Service Forestier 7509,
Sahamamy, Analamazaotra-Perinet RS, [18°56'S,
48°26'E], 21 Aug. 1953, bud (P, TEF); Service
Forestier 7564, Sahamaloto, Analamazaotra-Perinet
RS, [18°56'S, 48°26'E], 24 Jan. 1953, fl. (TEF);
Service Forestier 10348, Analamazaotra-Perinet RS,
[18°56'S, 48°26'E], 8 June 1954, bud (MO, P, TEF);
Service Forestier 14959, km 2, Perinet, Analamazaotra-
Perinet RS, [18°56'S, 48°26'E], 27 Feb. 1955, bud (P,
TEF); Service Forestier 17932, Analamazaotra-Perinet
RS, [18°56'S, 48°26'E], 800 m, 4 Apr. 1958, bud (P,
TEF); Service Forestier 21244, same locality, 26 July
1962, ster. (MO, P); Service Forestier 21278, 21279,
21280, 21281, 21282, same locality, 9 Nov. 1962,
ster. (MO, P); Service Forestier 21898, Sandrangato
FC, [19°06'30"S, 48°14'30"E], 900 m, 29 Oct. 1964,
bud (MO, P, TEF); Service Forestier 21955,
Sandrangato FC, pres Toby, PK 27 route d'Anosibe
An'ala, [19°06'30"S, 48°14'30"E], c. 900 m, 29 Oct.
1964, bud, fl., y.fr. (MO, P, TEF); Service Forestier
25441, Sandrangato FC, [19°06'30"S, 48°14'30"E],
900 m, 29 Oct. 1964, fl. (MO, P, TEF); Service
Forestier 26096, Sahatsaora (= Sahatsara), Zahamena
RNI, [17°38'30"S, 48°50'E], 19 July 1966, bud (P,
TEF); Service Forestier 26927, Nangaranana
(= Nagarana), Canton Marovoay, [18°44'S, 48°22'E],
Feb. 1970, bud (P, TEF); Service Forestier 26968,
same locality and date, fr. (P, TEF); Service Forestier
105-B-R-172, Ambodivoasary, Analamazaotra-Perinet
RS, [18°56'S, 48°26'E], 14 Mar. 1952, ster. (P, TEF);
Service Forestier 124-B-R-172, Menalamba, near
Analamazaotra-Perinet RS, [18°52'S, 48°22'30"E],
9 Apr. 1952, ster. (TEF); Thouvenot 12, Analamazaotra-
Perinet RS, [18°56'S, 48°26'E], 1919, bud, fl., y.fr.
(K, P); Viguier 851, Analamazaotra-Perinet RS,
[18°56'S, 48°26'E], 1000 m, 21 Oct. 1912, bud, fl,
y.fr. (MO, P).
Fig. 2. — Distributions of Leptolaena, mapped on the bioclima-
tic zones of Madagascar (after Cornet 1974; see Schatz 2000).
L. abrahamii (▲), L. delphinensis (★), L. multiflora (#).
2. Leptolaena cuspidata Baker
J. Linn. Soc. Bot. 25: 296 (1889). — Leptolaena
multiflora var. cuspidata (Baker) H. Perrier, Bull. Soc.
Bot. France 78: 60 (1931). — Type: Baron 5835 [cited
in protologue as "next 5836"], Madagascar, chiefly
from north-west Madagascar, fr. (holo-, K!; iso-, P!).
Leptolaena cuspidata is a shrub to small tree
that occurs in the subhumid forest of the
Sambirano region in NW Madagascar, below
500 m elevation (Fig. 3). It can be recognized by
its distinctive densely golden yellow granular
farinose involucre and lanceolate to ovate, long-
cuspidate leaves.
ADANSONIA, ser. 3 • 2001 • 23 (2)
177
Schatz G.E., Lowry II P.P. & Wolf A.-E.
Fig. 3. — Distributions of Leptolaena, mapped on the bioclima-
tic zones of Madagascar (after Cornet 1974; see Schatz 2000).
L. cuspidata (▲), L. pauciflora (•).
Vernacular names. — Zahana, Zahena,
Zana.
Conservation status. — With an extent of
occurrence < 20,000 km2, an area of occupancy
< 2,000 km2, and only three of 14 sub-popula-
tions encompassed within protected areas, L. cus-
pidata is assigned a preliminary status of
Vulnerable (VU).
Material examined. — Baron 5836, 6283, with-
out precise locality; Baum 333, Ambodimanga;
Birkinshaw 1, Antsatsaka; Bosser 20152, Ampasindava;
Cours 5674, Mt. Ambohiparaka; Decary 938, 1165,
2095, Maromandia; Decary 14775, 14794, Ambilobe;
Derleth 89, Manongarivo RS; Gentry 11870, 57-
58 km N of Ambanja; Hildebrandt 3306, Vavatobe;
Humbert 18752, Ambanja; Humbert 19041,
Ambodimagodro; Humbert 25580, SW of Ambilobe;
Humbert 32649, Ankarana RS; Humbert 32873,
Ambohipiraka; Perrier de la Bâthie 3021, Ambanja;
Perrier de la Bâthie 3023, Maromandia; Perrier de la
Bdthie 5340, Ambohipiraka; Perrier de la Bdthie
15677, bas Sambirano; Phillipson 2017, Beramanja;
Ravololonanahary 37, Ambato; Service Forestier 3078,
Irano (Irarona) to Beramanja; Service Forestier 3141,
3142, Andilamboay; Service Forestier 3171,
Tsaratanana RNI; Service Forestier 7501, Bekaka to
Benavony; Service Forestier 8225, Anjahakely
(= Ampahakely); Service Forestier 10434, Ambilobe;
Service Forestier 11486, Manongarivo RS; Service
Forestier 12987, Anjakely; Stiefel 68, Manongarivo RS;
Totozafy Be 563, Manongarivo RS.
3. Leptolaena delphinensis G.E. Schatz &
Lowry, sp. nov.
Haec species a congeneris ad Leptoaenam sensu stricto
pertinentibus foliis glabris apice obtusis usque rotundatis,
involucro urceolato granulari dense ferrugineo c. 3 mm
alto atque sepalis involucrum c. 1.5 mm excedentibus
distinguitur.
Typus. — McPherson, Dumetz & Rabevohitra
14222, Madagascar, Prov. Toliara, North of Fort
Dauphin near Ste. Luce (Manafiafy), NW of village in
forest remnant on sand, 24°47'S, 47°10'E, 20 m,
22 Oct. 1989, fl. (holo-, MO!; iso-, P!, TAN!, TEF!).
Shrubs to trees 3-10 m tall, trunk to 20 cm
in diam., stems glabrous. Leaves chartaceous,
narrowly to broadly elliptic, 1.8-5.2 X 0.8-
3.2 cm, reduced in the inflorescence, glabrous,
base obtuse to rounded, margins slightly to
strongly revolute, apex rounded to occasionally
obtuse, venation obscurely brochidodromous,
secondary veins (8-)11-13 per side, midrib
flat above, raised below; petiole 3 mm long,
c.1 mm diam., canaliculate, glabrous, stipules
indistinct, caducous. Inflorescences terminal,
corymbose, 10-50-flowered, the axes glabrous;
peduncle 0.7-0.8 mm long, involucre urceolate,
3 mm tall, 2.5 mm diam. at apex, densely gran-
ular ferruginous farinose, apical margin with 8-
9 subequal triangular teeth, 0.5 mm tall,
0.4 mm broad at base; sepals exserted 1-1.5 mm
beyond involucre, densely white hirsute outside,
178
ADANSONIA, ser. 3 • 2001 • 23 (2)
Revision of Leptolaena (Sarcolaenaceae)
Fig. 4. — Leptolaena delphinensis: A, flowering branch; B, flower (McPherson et al. 14222).
glabrous to sparsely hirsute inside; petals nar-
rowly elliptic, 7 X 2.5 mm, white fading to
yellow, exserted 6 mm beyond involucre at
anthesis; stamens 10, free to the base, filaments
c. 7-8 mm long, anthers 0.8 mm tall; ovary
conic-ovoid, densely whitish tomentose, style
slender, exserted, c. 7-8 mm long, stigma hemi-
spheric, with 3 weakly differentiated lobes,
c. 0.8 mm across. Fruit globose to widely
ovoid, somewhat asymmetric, with a longitudi-
nal channel and concave apex when dry, c. 3-
4 mm tall, 2-3 mm diam., glabrous, entirely
enclosed within the persistent sepals and
involucre. — Fig. 4.
Leptolaena delphinensis is a shrub to small tree,
and is restricted to humid littoral forest on sand
near Ft. Dauphin in SE Madagascar (Fig. 2). It
can be distinguished by its leaves with an obtuse
to rounded apex, small, urceolate, densely granu-
lar ferruginous involucre 3 mm tall, and sepals
that are exserted 1-1.5 mm beyond the involucre.
Vernacular names. — Fotobavy, Fotoda,
Fotondahy, Fonto, Fotona.
Conservation status. — With no sub-popu-
lations recorded from within the current pro-
tected areas network, and thus a predicted
population reduction of > 80% in the next three
ADANSONIA, ser. 3 • 2001 • 23 (2)
179
Schatz G.E., Lowry II P.P. & Wolf A.-E.
generations, L. delphinensis is assigned a prelimi-
nary status of Critically Endangered (CR). Along
with L. raymondii, the status of L. delphinensis
demonstrates the urgent need for additional pro-
tection of littoral forest along the East coast of
Madagascar.
Paratypes. — Madagascar, Prov. Toliara:
Decary 10064, Vinanibe, [25°03'S, 46°56'E],
10 July 1932, fl. (P); Dumetz et al. 683, Mandena
STF, 24°57'S, 47°00'E, 0-10 m, 17 Apr. 1989, bud
(K, MO, P, TAN, TEF); Humbert 5955, environs
de Fort Dauphin, sables entre le pic St. Louis et la
mer, [25°03'S, 47°00'E], 20 Sep.-6 Oct. 1928, fl.
(MO, P); Jacquemin 1149, Mandena STF, [24°57'S,
47°00'E], 8 Dec. 1972, fr. (MO, P); Johnson WII37,
Mandena STF, [24°57'S, 47°00'E], 16 July 1963,
bud (P); McPherson et al. 14142, Mandena STF,
low forest on sand, 24°57'S, 47°00'E, 25 m, 17 Oct.
1989, fl. (K, MO, P, TAN, TEF); McPherson &
Dumetz 14665, Mandena STF, 24°57'S, 47°00'E,
25 m, 7 Dec. 1989, fr. (MO, TAN); McPherson et
al. 14835, Ste. Luce (Manafiafy), 24°47'S, 47°10'E,
20 m, 18 Jan. 1990, fr. (MO, TAN); Rabenan-
toandro et al. 272, Mandena STF, 24°57'11"S,
47°00'16"E, 5 m, 27 Sep. 2000, fl. (G, K, MO, P,
TEF); Rabenantoandro et al. 308, Mandena STF,
24°55'20"S 47°02'23"E, 0-50 m, 4 Oct. 2000, fl.
(G, K, MO, P, TEF); Rabevohitra 2053, Ste. Luce
(Manafiafy), foret au N. du village, 24°46'S,
47°09'E, 10 m, 20 Oct. 1989, fl. (MO, P, TAN,
TEF); Rabevohitra & Rabenantoandro 3701,
Mandena STF, 24°57'03"S 47°00'03"E, 5 m,
4 Oct. 2000, fl. (G, K, MO, P, TEF); Service
Forestier 399, Mandena STF, foret littorale,
[24°57'S, 47°00'E], 16 Feb. 1949, fr. (P, TEF);
Service Forestier 8207, same locality, 24 July 1953,
fl. (P, TEF); Service Forestier 14597, same locality,
1 Sep. 1955, fl. (P); Service Forestier 28656, vestiges
forestiers, sur dunes anciennes, sur la route de
Ste. Luce, [24°47'S, 47°11'E], 11 Dec. 1968, fr. (P,
TEF); Zarucchi et al. 7504, Mandena STF, coastal
low forest on sand, 24°57'S, 47°02'E, 10 m, 26 May
1991, bud (K, MO, P, TAN).
4. Leptolaena gautieri G.E. Schatz & Lowry, sp.
nov.
Haec species a congeneris ad Leptoaenam sensu stricto
pertinentibus foliis ovatis vel late ovatis apice acutis
usque acuminatis secus costam subtus indumento mani-
feste vestitis, involucro granulari-farinoso dense ferrugi-
neo plerumque trichomatibus suberectis albidis
c. 0.5 mm longis vestito atque petalis 10.5-12 mm longis
distinguitur.
Typus. — Gautier 2920, Madagascar, Prov.
Antsiranana, Manongarivo RS, Bekolosy, 14°02'S,
48°18'E, 1240 m, 29 Mar. 1996, fl. (holo-, MO!; iso-,
G, P!, TAN!, TEF!).
Trees 3-15 m tall, stems initially hirsute-
tomentose, then glabrescent. Leaves chartaceous,
ovate to broadly ovate, 1.1-4.8 X 0.5-3.7 cm,
reduced in the inflorescence, blade glabrous, base
obtuse to rounded or subcordate, apex acute to
acuminate, the very tip rounded, occasionally
somewhat falcate, venation obscurely brochido-
dromous, secondary veins 9-14 per side, midrib
flat to slightly raised above, sparsely to moder-
ately hirsute-tomentose, raised below with
sparsely to moderately dense sericeous to hirsute
indumentum; petiole 2.5-4 mm long, 1 mm in
diam., canaliculate, moderately to densely hir-
sute, stipules indistinct, caducous. Inflorescences
terminal and axillary from the uppermost leaves,
corymbose, 7-30-flowered, the axes densely grey
hirsute-tomentose; peduncle 0.1-0.5 mm long,
indistinct; involucre urceolate, 3-3.5 mm tall,
2.5-3 mm diam. at apex, tapering evenly to a
cuneate base, densely granular ferruginous fari-
nose, usually also with sparse to moderately dense
whitish golden partially erect hairs to 0.5 mm,
especially towards the base, apical margin with 8-
10 subequal triangular teeth, 1 mm tall, 0.5-
1 mm broad at base; sepals exserted 1.5 mm
beyond involucre, densely whitish golden hirsute
outside, glabrous inside; petals narrowly oblong
to oblanceolate, 10.5-12 X 1.5-2 mm, white to
pale yellow, with evident venation upon drying,
exserted 7-9 mm beyond involucre teeth at
anthesis, the apex rounded; stamens 10-12, free
to the base, filaments 6-8 mm long, anthers 0.5-
0.9 mm tall; ovary ovoid, densely light brown
tomentose, style slender, exserted, 6-8 mm long,
stigma obtriangular to hemispheric, 1 mm in
diam. Fruit globose, somewhat asymmetric, with
a longitudinal channel and concave apex when
dry, c. 2.5 mm tall, 2.5 mm diam., glabrous,
entirely enclosed within the persistent sepals and
involucre.— Fig.5.
Leptolaena gautieri is a small to medium-sized
tree that occurs in mid-elevation subhumid forest
from 800 m to over 1,200 m (Fig. 6). It can be
180
ADANSONIA, ser. 3 • 2001 • 23 (2)
Revision of Leptolaena (Sarcolaenaceae)
Fig. 5. — Leptolaena gautieri: A, flowering branch; B, leaf (adaxial surface); C, leaf (abaxial surface); D, base of leaf (abaxial surface)
and petiole; E, flower. (A, Gautier2361; B-E, Schatz 3233).
ADANSONIA, ser. 3 • 2001 • 23 (2)
181
Schatz G.E., Lowry II P.P. & Wolf A.-E.
Fig. 6. — Distributions of Leptolaena, mapped on the bioclima-
tic zones of Madagascar (after Cornet 1974; see Schatz 2000).
L. masoalensis (▲), L. gautieri (•), L. raymondii (★).
recognized by its ovate to broadly ovate leaves
with an acute to acuminate apex and evident
indumentum on the midvein below, its densely
ferruginous granular farinose involucre that usu-
ally also has slightly erect whitish trichomes
0.5 mm long, and petals that are 10.5-12 mm
long.
Etymology. — The species epithet honors
Laurent Gautier, our friend and colleague at the
herbarium in Geneva, who has significantly
advanced our knowledge of the flora of the
Manongarivo Special Reserve.
Vernacular names. — Anjananjana, Zahana.
Conservation status. — With an area of
occupancy < 2,000 km2, and only four of eight
sub-populations encompassed within the
protected areas network, L. gautieri is assigned a
preliminary status of Vulnerable (VU).
Paratypes. — Madagascar, Prov. Antsiranana:
Gautier & Chatelain 2344, Manongarivo RS, Bekolosy,
vallon en amont de la chute, 14°02'S, 48°18'E,
1100 m, 15 June 1994, fl. (G, TAN); Gautier 2361,
Manongarivo RS, Bekolosy, 14°02'S, 48°18'E, 800 m,
15 June 1994, fl. (G, MO, P, TAN); Gautier 3510,
Manongarivo RS, Ambahatra, cours superieur, 13°59'S,
48°26'E, 1210 m, 8 Mar. 1999 (G, MO); Humbert &
Capuron 24147, Mt. Anjenabe, vallee inferieure de
l'Androranga, affluent de la Bemarivo aux environs
d'Antongondriha, [14°18'S, 49°45'E], 1130 m, 3-
7 Nov. 1950, fr. (P); Malcomber &Rakotomalala 2608,
Manongarvio RS, Bekolosy, 14°02'S, 48°18'E, 1000-
1200 m, 4 Dec. 1993, fr. (MO, P, TAN); McPherson &
van der Werff16359, Manongarivo RS, above the vil-
lage of Ambodisakoana, 14°05'S, 48°20'E, 1100 m,
16 Oct. 1994, fr. (MO, P, TAN); McPherson & van der
Werff16396, same locality, 1100-1200 m, 18 Oct.
1994, bud (MO, P, TAN); Rakotomalaza 48,
Manongarivo RS, Bekolosy, 14°04'S, 48°17'E, 1000-
1480 m, Apr. 1993, fl. (MO, P, TAN); Rakotozafy 336,
plateau de Beankany, Ambanja, 12 Dec. 1963, bud (P,
TAN); A. Randrianasolo 307, Marojejy RNI, 14°29'S,
49°49'E, 300-800 m, 21 Apr. 1993, fl. (MO, P, TAN);
A. Randrianasolo 325, same locality, 500-600 m,
22 Apr. 1993, fl. (MO, P, TAN); Reserves Naturelles
2237, Tsaratanana RNI, Ambavala, Marovato, Ambanja,
26 Sep. 1950, fr. (MO, P, TAN); Schatz 3233,
Manongarivo RS, Bekolosy, 14°06'S, 48°17'E,
1000 m, 28 Mar. 1991, fl. (K, MO, P, TAN). Prov.
Toamasina: Labat et al. 3068, Moramanga, Andasibe,
Foret de Maromizaha, 18°57'56"S, 48°27'34"E,
1020 m, 1 Apr. 1999, fr. (BR, G, K, P, MO, TAN);
McPherson 17467, Ambatovy, 18°51'12"S,
48°18'48"E, 1100 m, 26 Feb. 1998, bud (MO, P,
TAN); Perrier de la Bâthie 14887, Analamazaotra-
Perinet RS, [18°56'S, 48°26'E], Oct. 1920, fl. (P);
Rakotomalaza 1046, Ambatovy, 18°51'07"S,
48°18'47"E, 1100 m, 30 Jan. 1997, bud (MO, P,
TAN); Rakotomalaza 1279, same locality, 18°51'08"S,
48°18'59"E, 990 m, 25 Mar. 1997, bud (MO, P,
TAN); Rakotomalaza 1286, same locality, 18°51'28"S,
48°17'43"E, 26 Mar. 1997, bud (MO, P, TAN);
Rakotomalaza 1309, same locality, 18°51'10"S,
48°18'49"E, 1050 m, 13 May 1997, fl. (MO, P, TAN);
Randriambololona et al. 17, Mantady PN, 18°56'S,
48°26'E, 960 m, 27 Oct. 1997, fr. (MO, P, TAN);
Service Forestier 10341, Mangorokely, Anosibe,
Moramanga, [19°26'S, 48°12'E], 19 May 1954, fl. (P).
Prov. Toliara: Humbert 20704, Mt. Vohimavo au Nord
d'Ampasimena, bassin de la Manampanihy, [24°19'S,
47°08'E], 830 m, 27-28 Mar. 1947, fl. (MO, P).
182
ADANSONIA, ser. 3 • 2001 • 23 (2)
Revision of Leptolaena (Sarcolaenaceae)
Fig. 7.— Leptolaena masoalensis: A, flowering branch; B, leaf (adaxial surface); C, leaf (abaxial surface); D, flower (Rahajasoa et al. 790).
5. Leptolaena masoalensis G.E. Schatz &
Lowry, sp. nov.
Haec species a congeneris ad Leptoaenam sensu stricto
pertinentibus foliis glabris apice obtusis usque rotundatis,
involucro ellipsoideo plerumque glabro interdum sparse
albido-hirsuto c. 7 mm alto sepalos omnino includente
atque staminibus tantum 6 ad 8 distinguitur.
Typus. — Rahajasoa, Rabe & Bernard 790,
Madagascar, Prov. Antsiranana, Masoala PN, Réserve
Naturelle d'Ambato, 15°17'S, 50°21'E, 380 m, 8 Oct.
1994, fl. (holo-, MO!; iso-, K!, P!, TAN!).
Shrubs to 2 m tall; stems glabrous to sparsely
pubescent. Leaves subcoriaceous, elliptic to
slightly obovate, 2.4-4 X 1.1-1.7 cm, glabrous,
discolorous when dry, dark olive brown above,
lighter olive green below, base cuneate to acute,
margins strongly revolute, apex rounded, vena-
tion weakly brochidodromous, secondary veins
c.8 per side, midrib flat above, raised below with
very sparse scattered trichomes or glabrous; peti-
ole 2-3 mm long, 0.6-0.8 mm diam., shallowly
canaliculate, stipules caducous. Inflorescences
ADANSONIA, sér. 3 • 2001 • 23 (2)
183
Schatz G.E., Lowry II P.P. & Wolf A.-E.
axillary in the terminal leaves, 2-8-flowered, axes
sparsely to densely puberulous; peduncle 1-2 mm
long, distinct, involucre ellipsoid to oblong,
glabrous, 7 mm tall, 3 mm diam., apical margin
with 8 narrowly triangular, subequal teeth 1 mm
tall, 1 mm broad at base, glabrous or sometimes
sparsely short whitish hirsute; sepals completely
included within the involucre, 4 X 4 mm, circular,
strongly inrolled, densely whitish hirsute outside,
glabrous inside; petals narrowly oblong, 12 X
3 mm, clear yellow with evident venation upon
drying, glabrous, exserted 5-6 mm beyond involu-
cre; stamens 6-8, free to the base, filament 8-
10.5 mm long, anthers unknown; ovary ellipsoid,
1 mm diam., densely whitish golden hirsute-
strigose, the trichomes to 2 mm long, style
11.5 mm long, slender, exserted 4-5 mm above
involucre, stigma capitate, 1 mm in diam. Fruit
unknown. — Fig. 7.
Leptolaena masoalensis is known only from the
type collection made at Ambato, a hill in the
northeastern part of the Masoala Peninsula (Fig. 6)
well known for its large population of Nepenthes
masoalensis Schmid-Hollinger. At first glance,
L. masoalensis may be mistaken for L. delphinensis,
with which it shares leaves with an obtuse to
rounded apex. However, L. masoalensis can be
easily distinguished by its much larger, ellipsoid
to oblong involucre 7 mm tall, sepals that are
completely included within the involucre at
anthesis, and leaves with only 8 secondary veins
per side versus (8-)11-12 in L. delphinensis.
Conservation status. — With an area of oc-
cupancy < 100 km2, and only a single known pop-
ulation that appears to lie just outside the limit of
Masoala National Park, L. masoalensis is assigned a
preliminary status of Critically Endangered (CR).
6. Leptolaena multiflora Thouars
Hist. Veg. Isles Austral. Afriq.: 41, t. 11 (1805). —
Type: Thouars s.n., Madagascar, without precise local-
ity (P!).
As here recognized, Leptolaena multiflora
occurs primarily in coastal forest mostly on sand,
with a few populations recorded farther inland,
probably on similar substrates (Fig. 2). In the
past, this species has been broadly circumscribed
to include all of the new taxa described here, as
well as the Sambirano endemic L. cuspidata. As
defined here, L. multiflora can be distinguished
by its glabrous, elliptic, usually almost equilateral
leaves, the largest of which usually exceed 5.5 cm
in length and dry an olive green to brownish
color. In addition, the sepals at anthesis exceed
the involucre teeth by (0-8-)1-1.8 mm, a feature
that separates L. multiflora from L. abrahamii.
Vernacular names. — Amaninombi-
lahimena, Amaninombilahy, Amaninombilahy a
petites feuilles, Anjananjana, Birana, Dilatra,
Dilatry Lahy, Fotona, Fotona Beravina, Fotonala,
Hazoambo, Hazomasy, Helana, Jana,
Manilombilahy, Manina ombilahy, Sarifatra,
Tsilaitra a petites feuilles.
Conservation status. — Despite an extent
of occurrence and area of occupancy far in excess
of 20,000 km2 and 2,000 km2 respectively, with
only four of c. 19 sub-populations occurring in
protected areas, and thus indicating a possible
population reduction in the next three genera-
tions of > 50%, L. multiflora is assigned a prelim-
inary status of Endangered (EN).
MATERIAL EXAMINED. — Baron 2632, Bojer s.n.,
Forbes s.n., Louvel 6, Thouars s.n., without precise
locality; Andrianjafy 66, Rantabe; Bernard 372,
Masoala PN; Birkinshaw 354, Mananara RB; Boivin
s.n., Foulpointe; d'Alleizette 268, Nosy-Be(?); Decary
5198, Ifandana; Humblot 349, Nosive; Perrier de la
Bdthie 14186, Mahanoro; Rabe 157, Beankoraka;
Rabenantoandro 563, Mahatsara STF; Rabevohitra
3808, Mahabo; Rahajasoa 424, Masoala PN;
Raholivelo 37, Tampolo STF; A. Randrianasolo 292,
Ilaka-Est, 458, Tampolo STF; Reserves Naturelles
2746, 3403, 5514, 9113, 9124, 9310, Masoala RNI;
Richard s.n., Foulpointe; Richard 4, 40, Ste. Marie;
Schedl 135, Ambila-Lemaitso STF; Service Forestier
4-R-7, Antetezana STF; Service Forestier 89-R-303,
Manampano; Service Forestier 174-R-116,
Ampasinambo; Service Forestier 745-R-1, Ambila-
Lemaitso STF; Service Forestier 1371, Andatsakala;
Service Forestier 1616, Ambila-Lemaitso STF; Service
Forestier 2414, Soanierano Ivongo; Service Forestier
3680, Tohakandra; Service Forestier 3809, Misevo;
Service Forestier 4679, Antetezana STF; Service
Forestier 4813, Ifanirea; Service Forestier 5107, Berano;
Service Forestier 5677, Foulpointe; Service Forestier
6252, Rantabe; Service Forestier 7494, Fampotakely;
184
ADANSONIA, ser. 3 • 2001 • 23 (2)
Revision of Leptolaena (Sarcolaenaceae)
Service Forestier 7510, Antetezana STF; Service
Forestier 8901, Rantabe; Service Forestier 9501,
Ampangalana Atsimo; Service Forestier 10053, 10076,
Tampolo STF; Service Forestier 12401, Amporoforo;
Service Forestier 12929, Manombo STF; Service
Forestier 13906, Amporoforo; Service Forestier 14504,
Ifanirea; Service Forestier 15215, Tampolo STF; Service
Forestier 15302, Manombo RS; Service Forestier
15305, Berano; Service Forestier 15378, Andrazaha;
Service Forestier 15493, Analavory; Service Forestier
16118, Ambazato; Service Forestier 16215, Evato;
Service Forestier 16472, 17820, Tampolo STF; Service
Forestier 18082, Vohibola; Service Forestier 19173,
Tampolo STF; Service Forestier 19535, Ampangalana
Atsimo; Service Forestier 21217, Ambila-Lemaitso
STF; Service Forestier 21471, Ambohimary; Service
Forestier 21493, Manombo STF; Service Forestier
29717, Lakato; Service Forestier 29860, Tampolo STF;
Service Forestier 29892, Ambila-Lemaitso STF; Service
Forestier 32831 , 34550, Mahatsara; Service Forestier
34575, Sahamalaza; Service Forestier 35140,
Mahatsara; Ursch 116, Tampina.
7. Leptolaena pauciflora Baker
J. Linn. Soc. Bot. 20: 96, 97 (1883). — Lectotype
(here designated): Baron 1390, Madagascar, in forests
of the province of Imerina (K!; iso-, MO!,
P(3 sheets)!).
Leptolaena turbinata Baker, J. Linn. Soc. Bot. 20: 97
(1883). — Leptolaena pauciflora var. turbinata
(Baker) H. Perrier, Bull. Soc. Bot. France 78: 60
(1931).— Type: Baron 1560, Madagascar, East
coast (holo-, K!; iso-, P!).
Leptolaena parviflora Scott-Elliot, J. Linn. Soc. Bot.
29: 6 (1891). — Lectotype (here designated): Scott-
Elliot 2554, Madagascar, woods near Fort Dauphin
(K!; iso-, P(2 sheets)!).
Leptolaena rubella Scott-Elliot, J. Linn. Soc. Bot. 29: 7
(1891). — Leptolaena pauciflora var. rubella (Scott-
Elliot) H. Perrier, Bull. Soc. Bot. France 78: 60
(1931).— Type: Scott-Elliot 2369, Madagascar, woods
near Fort Dauphin (holo-, K!; iso-, P(2 sheets)!).
Leptolaena pauciflora is a small shrub to
medium-sized tree occurring from humid littoral
forest at sea level to subhumid and montane
woodland and thicket up to 2,300 m (Fig. 3).
Other than habit, it is remarkably uniform in
vegetative and reproductive characters through-
out its broad ecological range, and is easily recog-
nized by its small, broadly ovate to circular or
rhombic leaves usually not exceeding 2 cm in
length.
Of the two syntypes cited in the protologue of
L. pauciflora (Baron 1390 and Meller s.n.), the
former is here chosen as the lectotype. Syntypes
were also cited for L. parviflora (Scott-Elliot 2554
and Scott-Elliot 2713), the first of which is here
chosen as the lectotype.
Vernacular names. — Amaninambilahy,
Anjananjana, Anjananjanalahy, Fotina, Foto,
Fotona, Fotona Madinidravina, Fotondahy,
Hazomaritra, Laro, Madiorano, Milaliambo-
madinika, Taolandaro.
Conservation status. — With an extent of
occurrence far exceeding 20,000 km2, and c. 33
sub-populations, six of which are present in pro-
tected areas, L. pauciflora is considered not
threatened, and assigned a preliminary status of
Lower Risk (LR).
Material examined. — Baron 1390, 1560, 2624,
Meller s.n., without precise locality; Andrianjafy 11,
Andranofeno-Sud; Barnett 502, Mt. Ibity; Benoist 759,
Ambila-Lemaitso STF; Boiteau 2568, Mahialambo;
Bosser 7944, Tampoketsa d'Ankazobe; Bosser 9969,
Sahatsio; Bosser 14138, pic St. Louis; Bosser 15978,
Tampoketsa; Bosser 18772, col des Tapias; Cours
2981, Ambila-Lemaitso STF; Cours 4646, Didy;
Cremers 1624, 2020, Ambohitantely RS; Cremers
2311, Marosiky; Croat 29218, Antsirabe; Croat
29910, Itremo; Decary 4026, 4038, Fort-Dauphin;
Decary 6353, 6388, 6502, 6508, Ambila-Lemaitso
STF; Decary 7302, Ankazobe; Decary 7473,
Ambohitantely RS; Decary 10011, pic Saint Louis;
Decary 13063, 13226, Ambatofinandrahana; Decary
14023, 14036, Faliarivo; Decary 14927, Tampoketsa
d'Ankazobe; Decary 17214, Manankazo STF; Decary
17311, Ambositra; Decary 17333, Ambatofinan-
drahana; Decary 17403, Ambositra; Descoings 338,
Fort-Dauphin; Dorr 3842, col des Tapias; Edmondson
95-57, Mandena STF; Geay 7507, 7618, 7707, 7926,
8708 Mananjary; Gentry 11363, Ambila-Lemaitso
STF; Gentry11836, Ambohitantely RS; Gereau 3294,
Evatra; Gereau 3423, Manafiafy; Gereau 5690,
Anjozorobe; Harder 1536, Anjozorobe; Humbert
4640, col des Tapias; Humbert 5864, Enivaha;
Humbert 5899, pic St.Louis; Humbert 7111, col des
Tapias; Humbert 11090, Ambohitantely RS; Humbert
13835, Andohahela RNI; Humbert 14080, Imonty;
Humbert 14477, Faliarivo; Humbert 20623,
Ampasimena; Humbert 20702, Vohimavo; Humbert
28059, Ambatofinandrahana; Humbert 29938,
Itremo; Jacquemin 934, Ambavarano; Jacquemin
1158, Mandromodromotra; Johnson WII 33,
Mandena STF; Jongkind 828, 932, col des Tapias;
Keraudren 1062, pic Saint Louis; Keraudren 24894,
ADANSONIA, s<§r. 3 • 2001 • 23 (2)
185
Schatz G.E., Lowry II P.P. & Wolf A.-E.
24997, Fort Dauphin; Keraudren 25712, Ivato-
Ambatofinandrahana; Keraudren 26009, Itremo;
Labat 3010, col des Tapias; Leeuwenberg 14443,
c.20 km Ivato-Itremo; Leeuwenberg 14589, Ambila-
Lemaitso STF; Leeuwenberg 14648, Anjozorobe;
Lewis 728, Ambila-Lemaitso STF; Louvel 33bis,
Analamazaotra-Perinet RS; Lowry 4375, 4397,
Anjozorobe, 5215, Mandena STF; McPherson 14164,
Ste. Luce; McPherson 14278, Analalava (Manan-
tenina); Messmer 781, Itremo; Miller 8777,
Anjozorobe; Morat 3315, Ivato; Morat 4411,
Andohahela RNI; Morat 4512, Antsiatsiaka; Peltier
2159, Fiadanana; Peltier 2184, Anjoma; Perrier de la
Bdthie 3004, Andringitra RNI; Perrier de la Bdthie
3007, Ambatofinandrahana; Perrier de la Bdthie 3012,
Matatana; Perrier de la Bdthie 4488, Vatovavy; Perrier
de la Bdthie 5337, Manankazo STF; Perrier de la
Bdthie 5341, Ibity; Perrier de la Bdthie 5342,
Tamatave; Perrier de la Bdthie 13180, Tsitondroina;
Perrier de la Bdthie 13213, Beveromay; Perrier de la
Bdthie 13574, Ibity; Phillipson 1850, Ambila-Lemaitso
STF; Rabenantoandro 62, Itremo; Rabevohitra 3770,
Mahabo; Rakotoarisoa 22, 32, Ambatofinandrahana;
Rakotoarisoa 59, Faliarivo FC; Rakotozafy 2682,
Anjozorobe; Randriamampionona 331, 569, Andohahela
RNI; Randriamanantena 119, Andohahela RNI;
Randrianaivo 75, Ambohitantely RS; A. Randrianasolo
224, Ambila-Lemaitso STF; A. Randrianasolo 232,
Ivato; A. Randrianasolo 283, Manantantely;
A. Randrianasolo 347, Manafiafy; Rauh 1445, Fort-
Dauphin; Reserves Naturelles 13-RN-XI, Andohahela
RNI; Reserves Naturelles 168, 202, Ambohitantely RS;
Reserves Naturelles 706, Andringitra RNI; Reserves
Naturelles 1416, 1499, Ambila-Lemaitso STF; Reserves
Naturelles 6507, Andringitra RNI; Reserves Naturelles
6706, Ambohitantely RS; Reserves Naturelles 7166,
Andringitra RNI; Reserves Naturelles 11247,
Zahamena RNI; Reserves Naturelles 11691,
Andringitra RNI; Schatz 1320, Ambila-Lemaitso STF;
Schatz 2961, Anjozorobe; Schatz 3955, Ankafobe;
Schatz 3964, Itremo; Schedl 133, Ambila-Lemaitso
STF; Scott-Elliot 2369, 2554, 2713, 2713bis, 2753,
Fort-Dauphin; Service Forestier 3-R-3, Ambohitantely
RS; Service Forestier 54-R-10, Itremo; Service Forestier
232, Manandona; Service Forestier 1078, 1122, 1569,
Ambila-Lemaitso STF; Service Forestier 2155,
Fianarantsoa; Service Forestier 2708, Ampandrambato;
Service Forestier 3259, Ambila-Lemaitso STF; Service
Forestier 3267, Mahanoro; Service Forestier 3811,
Anosivelo; Service Forestier 4157, 4707, 4711, Ambila-
Lemaitso STF; Service Forestier 4829, Mangatsiatra;
Service Forestier 4887, 4908, Ambila-Lemaitso STF;
Service Forestier 5625, Vohipaho; Service Forestier
5631, Marohita; Service Forestier 5737, Ambila-
Lemaitso STF; Service Forestier 6077, Mandena STF;
Service Forestier 7035, Mananjary; Service Forestier
7258, Ambila-Lemaitso STF; Service Forestier 7292,
Misevo; Service Forestier 7419, Mandena STF; Service
Forestier 7598, Fierenana; Service Forestier 7771,
Mandromodromotra; Service Forestier 7820, Mandena
STF; Service Forestier 8416, Ambohitantely RS; Service
Forestier 9506, Ampangalana Atsimo; Service Forestier
9540, Marohita; Service Forestier 9592, Ambohitantely
RS; Service Forestier 9647, Ambila-Lemaitso STF;
Service Forestier 9668, Manantenina; Service Forestier
11542, Faliarivo; Service Forestier 12988, Vatovary
(=Vatovory); Service Forestier 13180, Ambila-Lemaitso
STF; Service Forestier 13472, Ankijana; Service
Forestier 13701 , Ampangalana Atsimo; Service Forestier
14047, Antanambao; Service Forestier 14661,
Nosy-Varika; Service Forestier 14670, Analalava;
Service Forestier 14763, Ankijana; Service Forestier
15156, Anjozorobe; Service Forestier 15806, Sahavoay;
Service Forestier 16113, Nosy-Varika; Service Forestier
16119, Ambazato; Service Forestier 16806, Manankazo
STF; Service Forestier 18361, Ambohitantely
RS; Service Forestier 18771, Manohilahy; Service
Forestier 19626, Iamboala; Service Forestier
19695, Ambohimanana; Service Forestier 19874,
Ambohitantely RS; Service Forestier 22456,
Andohahela RNI; Service Forestier 25366,
Ampasimpotsy STF; Service Forestier 32214, Ambila-
Lemaitso STF; Service Forestier 34202, Ambohitantely
RS; Zarucchi 7345, Anjozorobe; Zarucchi 7411,
Ambila-Lemaitso STF.
8. Leptolaena raymondii G.E. Schatz & Lowry,
sp. nov.
Haec species a congeneris ad Leptoaenam sensu stricto
pertinentibus foliis falcatis apice longe cuspidatis margin-
ibus valde revolutis subtus indumento persistente ex
trichomatibus brevibus adpressis etiam saepe trichomati-
bus suberectis > 1 mm longis constante distinguitur.
TYPUS. — Randrianasolo & Ranaivojaona 638,
Madagascar, Prov. Toamasina, Dist. Fenerive-Est,
Mahatsara STF, 17°38'S, 49°29'E, 5 m, 10 Nov.
1999, fl. (holo-, MO!; iso-, G!, K!, P!, TAN!).
Trees 5-10 m tall, to 20 cm dbh, stems short
sericeous-tomentellous. Leaves chartaceous,
ovate to broadly ovate, usually strongly falcate,
2-4.5 X 0.7-2.3 cm, glabrous above, appressed
short sericeous below often mixed with slightly
erect trichomes > 1 mm long, sometimes sub-
glabrescent, base strongly asymmetrical with one
side obtuse to rounded and the other side
cuneate, margins strongly revolute, apex long
cuspidate, the very tip rounded, venation
obscurely brochidodromous, secondary veins 8-
10 per side, midrib slightly raised above,
186
ADANSONIA, ser. 3 • 2001 • 23 (2)
Revision of Leptolaena (Sarcolaenaceae)
Fig. 8. — Leptolaena raymondii: A, flowering branch; B, leaf (abaxial surface); C, fruiting branch D, leaf (abaxial surface); E, immature
fruits (A-B, Service Forestier 34104; C-E, Service Forestier 34554).
glabrous, raised below, moderately densely
sericeous; petiole 2-4 mm long, 0.7-0.8 mm in
diam., canaliculate, densely sericeous, stipules
indistinct, caducous. Inflorescences terminal and
axillary from the uppermost leaves, corymbose,
9-20-flowered, axes densely gray hirsute-tomen-
tose; peduncle 0.5-1 mm long, indistinct to
somewhat distinct; involucre urceolate, base
acute to obtuse, 2.5-3.5 mm tall, 2.5-3 mm
diam. at apex, densely granular ferruginous fari-
nose and also with moderately dense white
appressed trichomes to 0.3 mm long, apical
margin with 8-10 unequal triangular to nar-
rowly triangular teeth, 1-1.3 mm tall, 0.3-1 mm
broad at base; sepals exserted 1 mm beyond
involucre, 3 X 3 mm, very widely obovate, apex
rounded, strongly inrolled, densely whitish hir-
sute outside, glabrous inside; petals narrowly
oblong to oblanceolate, 8-9 X 2-3 mm, white to
pale yellow, glabrous, exserted 4-5 mm beyond
involucre, with evident venation upon drying,
the apex rounded; stamens (10-)11(-12), free,
ADANSONIA, ser. 3 • 2001 • 23 (2)
187
Schatz G.E., Lowry II P.P. & Wolf A.-E.
filaments 4.5-5.5 mm long, anthers oblong 0.6-
0.9 X 0.3 mm; ovary obovoid, 1 X 0.8 mm,
densely whitish hirsute, style slender, 5-6 mm
long, exserted 4 mm above involucre, sparsely
whitish velutinous towards the base, stigma
capitate, 1.2-1.5 mm diam. Mature fruit
unknown.— Fig.8.
Leptolaena raymondii is a small to medium-
sized tree known only from Mahatsara Station
Forestiere (Fig. 6). It is the only species of
Leptolaena sensu stricto with persistent indumen-
tum throughout the lower surface of the leaf
blade, consisting of both short appressed tri-
chomes and often longer partially erect trichomes
greater than 1 mm long. The leaves of this species
are further characterized by their strongly revo-
lute margins.
Etymology. — The species epithet honors
our friend and colleague Raymond Rabevohitra,
Head of the FOFIFA herbarium (TEF), who has
contributed so much to our knowledge of
Madagascar's woody flora.
Vernacular names. — Amaninombilahy,
Anjananjana.
Conservation status. — With an area of
occupancy < 100 km2, and only a single known
population that is not included within the cur-
rent protected areas network, L. raymondii is
assigned a preliminary status of Critically
Endangered (CR). Along with L. delphinensis, the
status of L. raymondii demonstrates the urgent
need for additional protection of littoral forest
along the East coast of Madagascar.
PARATYPES. — MADAGASCAR, Prov. Toamasina:
Rabenantoandro et al. 562, Mahatsara STF,
17°38'13"S, 49°29'03"E, 2 m, 7 July 2001, bud
(G, K, MO, P, TAN); Razafindramora et al. 150,
same locality and date, bud (K, MO, P, TEF); Service
Forestier 32648, Marovovonana, Mahatsara STF,
[17°38'S, 49°29'E], 15 June 1985, bud (MO, TEF);
Service Forestier 34104, Mahatsara STF, [17°38'S,
49°29'E], 19 Oct. 1990, fl. (MO, TEF); Service
Forestier 34407, Andranotsara, Mahatsara STF,
[17°38'S, 49°29'E], 1 Oct. 1992, bud (G, K, MO, P,
TEF, WAG); Service Forestier 34554, point d'essai,
Mahatsara STF, [17°38'S, 49°29'E], 1 Dec. 1993,
bud, fr. (MO, TEF); Service Forestier 35141, same
locality, 30 Aug.-4 Sep. 1999, bud (MO, TEF).
Acknowledgements
We wish to thank K. SlKES, S. ANDRIAMBOLOLONERA
and J. RAHARIMAMPIONONA for specimen and data
management; A. JOUY and Roger LALA for the fine
illustrations; J. RABENANTOANDRO for help with field
work; R. GEREAU for assistance with the Latin diag-
noses; Ph. MORAT and his staff for hospitality
extended at the Laboratoire de Phanerogamie in Paris;
and the curators of several herbaria (K, TAN and
TEF) for giving us access to their specimens. Field
work was conducted under collaborative agreements
between the Missouri Botanical Garden and the Parc
Botanique et Zoologique de Tsimbazaza and the
Direction de la Recherche Forestiere et Piscicole,
FOFIFA, Antananarivo, Madagascar. We gratefully
acknowledge courtesies extended by the Government
of Madagascar (Direction Generale de la Gestion des
Ressources Forestieres) and by the Association
Nationale pour la Gestion des Aires Protegees. Special
thanks go to the Departement de Biologie et Ecologie
Vegetale, Universite d'Antananarivo for encourage-
ment and assistance. This research was conducted with
support from U.S. National Science Foundation
grants DEB-9024749 and DEB-9627072 and from
the John D. and Catherine T. MACARTHUR Foundation,
the Liz CLAIBORNE and Art ORTENBERG Foundation,
LWO Inc., and the National Geographic Society.
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189