Leptolaena pauciflora Baker, 1883

Schatz, George E., Lowry Ii, Porter P. & Wolf, Anne-Elizabeth, 2001, Endemic families of Madagascar. VII. A synoptic revision of Leptolaena Thouars sensu stricto (Sarcolaenaceae), Adansonia (3) 23 (2), pp. 171-189 : 185-189

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 throughout its broad ecological range, and is easily recognized 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, Milaliambomadinika, Taolandaro.

CONSERVATION STATUS.— With an extent of occurrence far exceeding 20,000 km 2, and c. 33 sub-populations, six of which are present in protected 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, Ambatofinandrahana; 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; Gentry 11836, 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, 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-Périnet RS; Lowry 4375, 4397, Anjozorobe, 5215, Mandena STF; McPherson 14164, Ste. Luce; McPherson 14278, Analalava (Manantenina); 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 Bâthie 3004, Andringitra RNI; Perrier de la Bâthie 3007, Ambatofinandrahana; Perrier de la Bâthie 3012, Matatana; Perrier de la Bâthie 4488, Vatovavy; Perrier de la Bâthie 5337, Manankazo STF; Perrier de la Bâthie 5341, Ibity; Perrier de la Bâthie 5342, Tamatave; Perrier de la Bâthie 13180, Tsitondroina; Perrier de la Bâthie 13213, Beveromay; Perrier de la Bâthie 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, I v a t o; A. Randrianasolo 283, M a n a n t a n t e l y; A. Randrianasolo 347, Manafiafy; Rauh 1445, Fort- Dauphin; Réserves Naturelles 13-RN-XI, Andohahela RNI; Réserves Naturelles 168, 202, Ambohitantely RS; Réserves Naturelles 706, Andringitra RNI; Réserves Naturelles 1416, 1499, Ambila-Lemaitso STF; Réserves Naturelles 6507, Andringitra RNI; Réserves Naturelles 6706, Ambohitantely RS; Réserves Naturelles 7166, Andringitra RNI; Réserves Naturelles 11247, Zahamena RNI; Réserves 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 S T F; Service Forestier 18361, A m b o h i t a n t e l y RS; Service Forestier 18771, Manohilahy; Service F o r e s t i e r 1 9 6 2 6, I a m b o a l a; S e r v i c e F o r e s t i e r 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 marginibus valde revolutis subtus indumento persistente ex trichomatibus brevibus adpressis etiam saepe trichomatibus suberectis> 1 mm longis constante distinguitur.

TYPUS. — Randrianasolo & Ranaivojaona 638, Madagascar, Prov. Toamasina, Dist. Fénérive-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 × 0.7-2.3 cm, glabrous above, appressed short sericeous below often mixed with slightly erect trichomes> 1 mm long, sometimes subglabrescent, 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,

Revision of Leptolaena (Sarcolaenaceae)

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-tomentose; 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 farinose and also with moderately dense white appressed trichomes to 0.3 mm long, apical margin with 8-10 unequal triangular to narrowly triangular teeth, 1-1.3 mm tall, 0.3-1 mm broad at base; sepals exserted 1 mm beyond involucre, 3 × 3 mm, very widely obovate, apex rounded, strongly inrolled, densely whitish hirsute outside, glabrous inside; petals narrowly oblong to oblanceolate, 8-9 × 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,

filaments 4.5-5.5 mm long, anthers oblong 0.6- Acknowledgements

0.9 × 0.3 mm; ovary obovoid, 1 × 0.8 mm, We wish to thank K. SIKES, S. ANDRIAMBOLOLONERA densely whitish hirsute, style slender, 5-6 mm and J. RAHARIMAMPIONONA for specimen and data long, exserted 4 mm above involucre, sparsely management; A. JOUY and Roger LALA for the fine whitish velutinous towards the base, stigma illustrations; J. RABENANTOANDRO for help with field capitate, 1.2-1.5 mm diam. Mature fruit work noses;; R Ph. G. EREAU MORAT for and assistance his staff with for the hospitality Latin diagunknown.— Fig. 8 . extended at the Laboratoire de Phanérogamie in Paris; and the curators of several herbaria (K, TAN and Leptolaena raymondii is a small to medium- TEF) for giving us access to their specimens. Field sized tree known only from Mahatsara Station work was conducted under collaborative agreements Forestière (Fig. 6). It is the only species of between Botanique the et Missouri Zoologique Botanical de Tsimbazaza Garden and the and Parc the

Leptolaena sensu stricto with persistent indumen- Direction de la Recherche Forestière et Piscicole, tum throughout the lower surface of the leaf FOFIFA, Antananarivo, Madagascar. We gratefully blade, consisting of both short appressed tri- acknowledge courtesies extended by the Government chomes and often longer partially erect trichomes of Madagascar (Direction Générale de la Gestion des Ressources greater than 1 mm long. The leaves of this species Nationale pour Forestières la Gestion) des and Aires by the Protégées Association. Special are further characterized by their strongly revo- thanks go to the Département de Biologie et Écologie lute margins. Végétale, Université d’Antananarivo for encouragement and assistance. This research was conducted with ETYMOLOGY.— The species epithet honors support from U.S. National Science Foundation grants DEB-9024749 and DEB-9627072 and from our friend and colleague Raymond RABEVOHITRA, the John D. and Catherine T. MACARTHUR Foundation, Head of the FOFIFA herbarium ( TEF), who has the Liz CLAIBORNE and Art ORTENBERG Foundation, contributed so much to our knowledge of LWO Inc., and the National Geographic Society.

Madagascar’s woody flora.

VERNACULAR NAMES.— Amaninombilahy,

Anjananjana. REFERENCES

CONSERVATION STATUS.— With an area of

occupancy <100 km 2, and only a single known ALVERSON W.S., KAROL K.G., BAUM D.A, CHASE population that is not included within the cur- M.W., SWENSEN S.M., MCCOURT R. & SYTSMA rent protected areas network, L. raymondii is K.J. 1998. — Circumscription of the Malvales and assigned a preliminary status of Critically relationships to other Rosidae: evidence from rbc L Endangered ( CR). Along with L. delphinensis , the sequence data. Amer. J. Bot. 85: 876-887.

BAYER C., FAY M.F., DE BRUIJN A.Y., SAVOLAINEN V., status of L. raymondii demonstrates the urgent MORTON C.M., KUBITZKI K., ALVERSON W.S. & need for additional protection of littoral forest CHASE M.W. 1999. — Support for an expanded along the East coast of Madagascar. family concept of Malvaceae within a recircumscribed order Malvales : a combined analysis of plastid atp B and rbc L DNA sequences. Bot. J. Linn. PARATYPES. — MADAGASCAR, Prov. Toamasina: Soc. 129: 267-303.

Rabenantoandro et al. 562, Mahatsara STF, CAPURON R. 1970. — Observations sur les Sarco- 17°38’13”S, 49°29’03”E, 2 m, 7 July 2001, bud laenacées. Adansonia , sér. 2, 10: 247-265.

(G, K, MO, P, TAN); Razafindramora et al. 150, CAVACO A. 1951. — Remarques sur les genres same locality and date, bud (K, MO, P, TEF); Service Leptolaena et Xerochlamys (Chlaenacées) . Un nou- Forestier 32648, Marovovonana, Mahatsara STF, veau genre de Chlaenaceae . Bull. Mus. Natl. Hist. [17°38’S, 49°29’E], 15 June 1985, bud ( MO, TEF); Nat ., sér. 2, 23: 133-139.

Service Forestier 34104, Mahatsara STF, [17°38’S, CAVACO A. 1952a. — Recherches sur les Chlénacées, 49°29’E], 19 Oct. 1990, fl. ( MO, TEF); Service famille endémique de Madagascar. Mém. Inst. Sci. Forestier 34407, Andranotsara, Mahatsara STF, Madagascar, sér. B, Biol. Vég. 4: 59-92 .

[17°38’S, 49°29’E], 1 Oct. 1992, bud (G, K, MO, P, CAVACO A. 1952b. — Chlénacées. Fl. Madagascar TEF, WAG); Service Forestier 34554, point d’essai, 126: 1-37 .

Mahatsara STF, [17°38’S, 49°29’E], 1 Dec. 1993, COETZEE J.A. & MULLER J. 1984. — The phytogeobud, fr. ( MO, TEF); Service Forestier 35141, same graphic significance of some extinct Gondwanan locality, 30 Aug.-4 Sep. 1999, bud ( MO, TEF). pollen types from the Tertiary of the southwestern Cape ( South Africa). Ann. Missouri Bot. Gard. 71: SCHATZ G.E. 2000. — Endemism in the Malagasy 1088-1099. tree flora: 1-9, in LOURENÇO W.R. & GOODMAN CORNET A. 1974. — Essai de cartographie bioclima- S.M. (eds.), Diversity and Endemism in Madagascar. tique à Madagascar. Notic. Explic. ORSTOM No. 55. Mém. Soc. Biogéogr., Paris .

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Manuscript received 18 September 2001; revised version accepted 2 October 2001.

Figures

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).