Rhodolaena humblotii Baill.

Schatz, George E., Lowry Ii, Porter P. & Wolf, Anne-Elizabeth, 2000, Endemic families of Madagascar. VI. A synoptic revision of Rhodolaena (Sarcolaenaceae), Adansonia (3) 22 (2), pp. 239-252 : 246-252

5. Rhodolaena humblotii Baill.

Bull. Mens. Soc. Linn. Paris 1: 566 (1886). — Lectotype (here designated): Humblot 474, Madagascar, Antsianaka ( P!; iso-, K!, P!)

Rhodolaena humblotii is a medium-sized tree that occurs in low to mid-elevation humid to subhumid forest from Ifanadiana to Fampanambo NW of Maroantsetra ( Fig. 4 ). It can usually be recognized easily by its twigs with dense, long, golden indument, and generally persistent, linear to narrowly triangular stipules. Rhodolaena humblotii has recently been collected at several localities, including Betampona RNI, Zahamena PN/RNI, the upper Rantabe water- shed in the Bay of Antongil area, and at Fampanambo near Maroantsetra.

When BAILLON (1886 a) described R. humblotii , he cited two collections (Humblot 428 and 474), the latter of which is in better condition and has therefore been selected as the lectotype.

VERNACULAR NAMES.— Arinala, Hazomafana,

Fotona, Malemisisika, Vandroazana, Voandrozana.

MATERIAL EXAMINED. — Andrianjafy 68, Beanana, 88, Betampona RNI, 102, Zahamena RNI; Cours 1844 (= Herb. Inst. Sci. Madag. 1844), Ambodivoangy, 2392, Sahalampy, 2429 (= Herb. Inst. Sci. Madag. 2429 and Herb. St. Agric. Alaotra 2429), Sahalampy à Ampitanonoka; Guillaumet 2420, Antanambao-Manampotsy; Herb. Inst. Sci. Madag. 3723, Zahamena RNI; Homolle 1844, without precise locality; Humblot s.n., without precise locality, 428, 474, Antsianaka; Iambana 146, Betampona RNI; Jacquemin 226, Fotsialanana à Mitanonoka; Malcomber 2513, Zahamena RNI; Nicoll 501, Manambato; Pierre s.n., without precise locality; Rakotozafy 677, Fotsialanana-Ampitanonoka; Ratovoson 92, Fampanambo; Réserves Naturelles 3723, Zahamena RNI, 7414, Betampona RNI, 7834, Zahamena RNI, 8749, Betampona RNI, 10508, 12442, Zahamena RNI; Service Forestier 4484, Sahamazaka, 9060, Beanana, 9109, Sahajinja- Sahamalaza, 17558, Betampona RNI, 21420, Tsinjoarivo, 23229, Ifanadiana, 31934, Verezanantsoro, Mananara-Nord PN.

6. Rhodolaena leroyana G.E. Schatz, Lowry & A.-E. Wolf, sp. nov.

Haec species a congeneris foliorum apicibus acutis acuminatisve munitis venatione secundaria rectangulari, ab omnibus involucro maturitate profunde laciniato atque seminibus indumento albo longo densoque obtectis distinguitur.

TYPUS. — Réserves Naturelles 124, Madagascar, Prov. Toamasina, Canton Ambodiriana, Anjiro à la RN1 [Betampona], sommet, [17°55’S, 49°13’E], 19 Mar. 1954, fl. (holo-, P!; iso-, K!, MO!, P!, TEF!) .

Shrub to tree to 8 m tall, branches glabrous. Leaves with petiole 0.4-0.7 cm long, lamina membranous to subcoriaceous, glabrous, elliptic to slightly obovate, 3.6-12 × 1.3-4.7 cm, base acute to cuneate, apex acuminate to cuspidate, acumen 0.3-1.3 cm long, venation weakly brochidodromous with 5-6 (-11) secondary veins per side nearly perpendicular to the midvein, midvein slightly sunken above, prominently raised below. Inflorescence solitary, axillary, 2-flowered, peduncle slender, 3.5-8 cm long, involucre 5-lobed, with dense golden simple and stellate indument, pedicel 0.7 cm long; outer 2 sepals narrowly elliptic, 0.2-0.7 × 0.2 cm, base attenuate and abruptly truncate, apex acute, the lower half strongly reflexed and then the upper half abruptly bent upwards, the margins revolute for the lower half and then involute for the upper half, densely stellate pubescent outside, glabrous inside; inner 3 sepals ovate, 1.7-2.5 × 1.2-1.4 cm, strongly concave, base truncate, apex obtuse to acute, densely stellate pubescent outside, glabrous inside; petals elliptic, 3.8-4 × 1.7 cm, base Prov. Antsiranana, massif de Tsihomanaomby, ca.

cuneate and abruptly truncate, apex obtuse to 35 km au Nord de Sambava, W d’Ambavala, forêt rounded; disc annular, 0.4 mm in diam., 0.2 mm 50 dense °02’52 sur ”E pente, 150 m, sur, 7 May sable 2000 quartzite, fr. (holo-, 14, ° MO 06’03!; iso- ”S,,

high; stamens ca. 40, inserted above disc at base K!, P!, TAN!, TEF!).

of ovary, 3.5 cm long, anthers basifixed and

resupinate, 1 × 1 mm; ovary subglobose, 4 mm Tree to 10 m tall, branches glabrous. Leaves high, 4 mm in diam., tomentose, style to 5 cm with petiole 0.7-1.6 cm long, canaliculate, long, exerted, stigma crateriform. Fruit 3-valved, glabrous; lamina stiff coriaceous, drying olive ultimately completely surrounded by the some- green, glossy, narrowly elliptic to elliptic to what fleshy, accrescent involucre with deeply slightly obovate-elliptic, 3.2-11.2 × 2-6.8 cm, laciniate margins, valves broadly ovate, 1.3 cm base acute to cuneate and slightly asymmetrical, long, 1.5-1.7 cm broad, densely golden pubes- apex abruptly short acuminate, occasionally cent outside; seeds depressed globose, 1.2 mm in rounded, margin weakly to strongly revolute, diam., 0.5 mm thick, with dense white wooly venation brochidodromous with 9-11 secondary indument.— Fig. 5 . veins per side strongly ascending at 45º angle or less, the secondary veins only slightly raised on Rhodolaena leroyana is a shrub to small tree in upper surface; flowers known only at post-anthelow elevation humid forest, and is known only sis; peduncles in fruit 4.5-7.5 cm long, 2 mm in from the Betampona RNI, where it was collected diam.; involucre entire, shallowly 3-lobed, lobes earlier this year ( Fig. 1 ). It is easily distinguished 3 mm long, 5 mm broad, densely light olive-tan by its leaves with an acuminate apex and 2º veins stellate pubescent; pedicel post-anthesis 3-4 mm nearly perpendicular to the midvein (> 80º), as long, 2-3 mm in diam., expanding in fruit to well as its densely golden tomentose ovary and 5-7 mm long, 4 mm in diam.; outer 2 sepals trifruit, and seeds covered with a dense, long, white angular to ovate, 1-2 × 0.6-2 mm, inner 3 sepals indument. broadly elliptic, 2.2-2.7 × 1.3-1.8 cm; stamens ca. 60, filament 4.5-6 cm long, anther 0.7 mm ETYMOLOGY.— The species epithet honors the long; style 6.5 cm long, stigma capitate with late Jean-François LEROY. 3 triangular lobes; fruits globose, 1.5-1.7 cm in diam., weakly 3-sided in cross section, valves of V E R N AC U L A R N A M E S.— Manasavelona, dehisced fruits ovate, 2-3.3 cm long, 2-2.5 cm Tsipatika, Tsipatikala. broad, densely stellate-strigose, with the central trichome branch much longer than the others.— P

ARATYPES

. — M

ADAGASCAR

, Prov. Toamasina:

Andrianarisata 127, Betampona RNI, 17°55’S, Fig. 2 .

49°13’E, 24 Apr. 1994, y.fr. ( MO, P, TEF), 171 ,

Betampona RNI, 17°55’S, 49°13’E, 29 Apr. 1994, Rhodolaena macrocarpa is a medium-sized tree y.fr. ( MO, P, TEF), 224, Betampona RNI, 17°55’S , that occurs in low elevation humid forest, and is 49°13’E, 5 Oct. 1994, fr. ( MO!, P, TEF); Andrianjafy known only from the type locality N of Sambava 86, Betampona RNI, 17°55’50”S, 49°12’12”E , 550

m, 17 July 2000, fr. (K, MO, P, TAN, TEF); Service ( Fig. 4 ). It is distinguished by its stiff coriaceous Forestier 3573, Betampona RNI, [17°55’S, 49°13’E], leaves drying glossy olive green with secondary 12 June 1951, fr. ( P, TEF). veins strongly ascending at a 45º angle or less to the midvein, and its large fruit, exceeding those

of all other members of the genus. Based on the 7. Rhodolaena macrocarpa G.E. Schatz, Lowry available material, R. macrocarpa appears to be & A.-E. Wolf, sp. nov. the only member of the genus with flowers borne Haec species a congeneris foliorum venatione secun- singly above the involucre, whereas in the other daria divergente, floribus solitariis atque fructum magni species they consistently occur in pairs, one of valvis 2-3.3 × 2-2.5 cm distinguitur. which develops into fruit and the other aborts, TYPUS. — Andrianjafy, Rabenantoandro, leaving a more or less evident scar.

Razafindrakoto, Birkinshaw & Lowry 61, Madagascar, VERNACULAR NAME.— Voandrozana.

Fig. 6. — Photographs of Rhodolaena . — R. bakeriana : A, flowering branch; B, flower. — R. acutifolia : C, flowering branch. — R. leroyana : D, immature fruit. — R. macrocarpa : E, fruiting branch; F, immature fruit (Photos – A, G.E. SCHATZ; B, J.-N. LABAT; C, S. MALCOMBER; D, M. ANDRIANJAFY; E-F, P. P. LOWRY).

PARATYPES. — MADAGASCAR, Prov. Antsiranana: scribed order Malvales : a combined analysis of plas- Service Forestier (Capuron) 27200, Madagascar, Prov. tid atp B and rbc L DNA sequences. Bot. J. Linn. Soc. Antsiranana, entre Andrangana et la rivière 129: 267-303 .

Anjambazamba (route de Sambava à Antsirabe-Nord), CAPURON R. 1970. — Observations sur les [14°05’S, 50°03’E], 2-7 Dec. 1966, fr. (K, MO, P, Sarcolaenacées. Adansonia , sér. 2, 10: 247-265.

TEF). CAVACO A. 1952a. — Recherches sur les Chlénacées,

famille endémique de Madagascar. Mém. Inst. Sci.

Madagascar, sér. B, Biol. Vég. 4: 59-92.

CAVACO A. 1952 b. — Chlénacées. Fl. Madagascar 126: Acknowledgements 1-37.

We wish to thank C. BIRKINSHAW, K. SIKES, COETZEE J.A. & MULLER J. 1984. — The phytogeo- S.A NDRIAMBOLOLONERA, J. RAHARIMAMPIONONA and graphic significance of some extinct Gondwanan L. ANDRIAMAHEFARIVO for specimen and data man- pollen types from the Tertiary of the southwestern agement; M. ANDRIANJAFY for helpful comments, field Cape (South Africa). Ann. Missouri Bot. Gard. 71: observations and several recent collections; A. JOUY for 1088-1099.

the fine illustrations; R. GEREAU for assistance with the CORNET A. 1974. — Essai de cartographie bioclima- Latin diagnoses; and Ph. MORAT and his staff for hos- tique à Madagascar. Notic. Explic. ORSTOM pitality extended at the Laboratoire de Phanérogamie No.55.

in Paris. Field work was conducted under collaborative DU PUY D.J. & MOAT J. 1996. — A refined classificaagreements between the Missouri Botanical Garden tion of the vegetation types of Madagascar, and and the Parc Botanique et Zoologique de Tsimbazaza their current distribution: 205-218, in LOURENÇO and the Direction de la Recherche Forestière et W.R. (ed.), Biogéographie de Madagascar. Editions Piscicole, FOFIFA, Antananarivo, Madagascar. We de l’ORSTOM, Paris.

gratefully acknowledge courtesies extended by the LOWRY II P. P., SCHATZ G.E. & PHILLIPSON P.B. Government of Madagascar (Direction Générale de la

1997.— The classification of natural and anthro- Gestion des Ressources Forestières) and by the pogenic vegetation in Madagascar: 93 -123, in Association Nationale pour la Gestion des Aires GOODMAN S.M. & PATTERSON B.D. (eds.), Protégées. This research was conducted with support Natural change and human impact in Madagascar. from U.S. National Science Foundation grants Smithsonian Inst. Press, Washington & London. DEB-9024749 and DEB-9627072 and from the LOWRY II P. P., SCHATZ G.E. & PHILLIPSON P.B. John D. and Catherine T. MACARTHUR Foundation, 1998.— Classification de la végétation naturelle et the Liz CLAIBORNE and Art ORTENBERG Foundation, anthropique de Madagascar. Bull. Acad. Natl. LWO Inc., and the National Geographic Society. Madagascar, n.s., 73: 219-233.

LOWRY II P. P., SCHATZ G.E., LEROY J.-F. & WOLF A.-E.

1999. — Endemic families of Madagascar. III. A

synoptic revision of Schizolaena (Sarcolaenaceae) . REFERENCES Adansonia , sér. 3, 21: 183-212.

LOWRY II P. P., HAEVERMANS T., LABAT J.-N.,

SCHATZ G.E., LEROY J.-F. & WOLF A.-E. 2000. — ALVERSON W.S., KAROL K.G., BAUM D. A, CHASE M.W., Endemic families of Madagascar. V. A synoptic SWENSEN S.M., MCCOURT R. & SYTSMA K.J. revision of Eremolaena, Pentachlaena, and 1998.— Circumscription of the Malvales and rela- Perrierodendron (Sarcolaenaceae). Adansonia, sér. 3, tionships to other Rosidae: evidence from rbc L 22: 11-31.

sequence data. Amer. J. Bot. 85: 876-877. SCHATZ G.E. 2000. — Endemism in the Malagasy BAILLON H. 1886a. — Liste des plantes de tree flora: 1 -9, in LOURENÇO W.R. & GOODMAN S.M. Madagascar (suite). Bull. Mens. Soc. Linn. Paris 1: (eds.), Diversity and Endemism in Madagascar. 562-568. Mém. Soc. Biogéogr., Paris.

BAILLON H. 1886 b. — Nouvelles observations sur SCHATZ G.E., BIRKINSHAW, C.R., LOWRY II P. P., les Chlénacées. Bull. Mens. Soc. Linn. Paris 1: RANDRIANTAFIKA F. & RATOVOSON F. 2000. — 570-572. The endemic plant families of Madagascar project: BAKER J.G. 1883. — Contributions to the flora of integrating taxonomy and conservation: 11-24, in Madagascar. Part I. Polypetalae. J. Linn. Soc. Bot. LOURENÇO W.R. & GOODMAN S.M. (eds.), 20: 87-158. Diversity and Endemism in Madagascar. Mém. Soc. BAKER J.G. 1886. — Further contributions to the Biogéogr., Paris.

flora of Madagascar. J. Linn. Soc. Bot. 22: 441-537. SCHATZ G.E., LOWRY II P. P., LESCOT M., WOLF A.-E., BAYER C., FAY M.F., DE BRUIJN A.Y., SAVOLAINEN V., ANDRIAMBOLOLONERA S., RAHARIMALALA V. & MORTON C.M., KUBITZKI K., ALVERSON W.S. & RAHARIMAMPIONONA J. 1996. — Conspectus of the CHASE M.W. 1999. — Support for an expanded vascular plants of Madagascar: a taxonomic and family concept of Malvaceae within a recircum- conservation electronic database: 10 -17, in VAN DER

MAESEN L.J.G., VAN DER BURGT X.M. & VAN MEDENBACH DE ROOY J.M. (eds.), The Biodiversity of African Plants. Proc. XIV AETFAT Congress. Klower Academic Publishers, Wageningen, The Netherlands.

SCHATZ G.E., LOWRY II P. P. & WOLF A.-E. 1998. — Endemic families of Madagascar. I. A synoptic revision of Melanophylla (Melanophyllaceae). Adansonia, sér. 3, 20: 233-242.

SCHATZ G.E., LOWRY II P. P. & WOLF A.-E. 1999 a. — Endemic families of Madagascar. II. A synoptic revision of Sphaerosepalaceae. Adansonia, sér. 3, 21: 107-123.

SCHATZ G.E., LOWRY II P. P. & WOLF A.-E. 1999 b. — Endemic families of Madagascar. IV. A synoptic revision of Asteropeia (Asteropeiaceae). Adansonia, sér. 3, 21: 255-268.

Manuscript received 29 May 2000;

revised version accepted 27 July 2000.

Figures

Fig. 4. — Distributions of Rhodolaena, mapped on the bioclimatic zones of Madagascar (after CORNET 1974; see SCHATZ 2000). R. humblotii (•), R. macrocarpa (*).

Fig. 5. — Rhodolaena leroyana: A, flowering branch; B, fruiting branch; C, leaf (abaxial surface); D, inflorescence (post-anthesis); E, ovary; F, anther; G, sepal; H, petal; J, seed. (A, C-H, Service Forestier 3573; B, J, Andrianarisata 224).

Fig. 1. — Distributions of Rhodolaena, mapped on the bioclimatic zones of Madagascar (after CORNET 1974; see SCHATZ 2000). R. acutifolia (•), R. altivola (*), R. bakeriana (Δ), R. leroyana (-).

Fig. 2. — Rhodolaena coriacea: A, flowering branch; B, leaf (adaxial surface); C, leaf (abaxial surface); D, flower (schematic view); E, sepals (inner, outer). — Rhodolaena macrocarpa: F, fruiting branch; G, leaf (adaxial surface); H, leaf (abaxial surface); J, dehisced fruit with involucre (A, Service Forestier 8893; B-E, Service Forestier 8864; F, Andrianjafy et al. 61; G-J, Service Forestier 27200).