Leptolaena cuspidata Baker, 1889

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 : 177-185

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, longcuspidate leaves.

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 Bâthie 5340, Ambohipiraka; Perrier de la Bâthie 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!) .

Fig. 3. — Distributions of Leptolaena , mapped on the bioclima- Shrubs to trees 3-10 m tall, trunk to 20 cm tic zones of Madagascar (after CORNET 1974; see SCHATZ 2000). L. cuspidata (Δ), L. pauciflora (•). in diam., stems glabrous. Leaves chartaceous,

narrowly to broadly elliptic, 1.8-5.2 × 0.8-

3.2 cm, reduced in the inflorescence, glabrous,

VERNACULAR NAMES.— Zahana, Zahéna, base obtuse to rounded, margins slightly to Zana. strongly revolute, apex rounded to occasionally CONSERVATION STATUS.— With an extent of obtuse, venation obscurely brochidodromous, occurrence <20,000 km 2, an area of occupancy secondary veins (8-)11-13 per side, midrib <2,000 km 2, and only three of 14 sub-popula- flat above, raised below; petiole 3 mm long, tions encompassed within protected areas, L. cus- c. 1 mm diam., canaliculate, glabrous, stipules pidata is assigned a preliminary status of indistinct, caducous. Inflorescences terminal, Vulnerable (VU). corymbose, 10-50-flowered, the axes glabrous;

peduncle 0.7-0.8 mm long, involucre urceolate,

MATERIAL EXAMINED. — Baron 5836, 6283, with- 3 mm tall, 2.5 mm diam. at apex, densely granout precise locality; Baum 333, Ambodimanga; ular ferruginous farinose, apical margin with 8- Birkinshaw 1, Antsatsaka; Bosser 20152, Ampasindava; Cours 5674, Mt. Ambohiparaka; Decary 938, 1165, 9 subequal triangular teeth, 0.5 mm tall, 2095, Maromandia; Decary 14775, 14794, Ambilobe; 0.4 mm broad at base; sepals exserted 1-1.5 mm Derleth 89, Manongarivo RS; Gentry 11870, 57- beyond involucre, densely white hirsute outside,

glabrous to sparsely hirsute inside; petals narrowly elliptic, 7 × 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 hemispheric, with 3 weakly differentiated lobes, c. 0.8 mm across. Fruit globose to widely ovoid, somewhat asymmetric, with a longitudinal 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 granular 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-populations recorded from within the current protected areas network, and thus a predicted population reduction of> 80% in the next three

generations, L. delphinensis is assigned a preliminary status of Critically Endangered (CR). Along with L. raymondii , the status of L. delphinensis demonstrates the urgent need for additional protection 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) ; Rabenantoandro 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), forêt 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, M a n d e n a S T F, f o r ê t l i t t o r a l e, [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 manifeste vestitis, involucro granulari-farinoso dense ferrugineo 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 hirsutetomentose, then glabrescent. Leaves chartaceous, ovate to broadly ovate, 1.1-4.8 × 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 brochidodromous, secondary veins 9-14 per side, midrib flat to slightly raised above, sparsely to moderately 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 hirsute, 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 farinose, 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 × 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

CONSERVATION STATUS.— With an area of occupancy <2,000 km 2, 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 supérieur, 13°59’S,

48°26 ’ E, 1210 m, 8 Mar. 1999 (G, MO); Humbert &

Capuron 24147, Mt. Anjenabe, vallée inférieure 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 Werff 16359, 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

Werff 16396, 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); Réserves 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. Fig. 6. — Distributions of Leptolaena, mapped on the bioclima- Toamasina: Labat et al. 3068, Moramanga, Andasibe, tic zones of Madagascar (after CORNET 1974; see SCHATZ 2000). Forêt de Maromizaha, 18°57’56”S, 48°27’34”E, L. masoalensis (Δ), L. gautieri (•), L. raymondii (*). 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, recognized by its ovate to broadly ovate leaves TAN); Perrier de la Bâthie 14887, Analamazaotrawith an acute to acuminate apex and evident Périnet

Rakotomalaza RS, [18°56 1046 ’S,, 48 Ambatovy °26’E], Oct,. 18 1920 ° 51, ’ fl 07. (” P S);,

indumentum on the midvein below, its densely 48°18’47”E, 1100 m, 30 Jan. 1997, bud ( MO, P, ferruginous granular farinose involucre that usu- TAN); Rakotomalaza 1279, same locality, 18°51’08”S , ally also has slightly erect whitish trichomes 48°18’59”E, 990 m, 25 Mar. 1997, bud ( MO, P, 0.5 mm long, and petals that are 10.5-12 mm TAN); Rakotomalaza 1286, same locality, 18°51’28”S ,

48°17’43”E, 26 Mar. 1997, bud ( MO, P, TAN); long. Rakotomalaza 1309, same locality , 18°51’10”S,

48°18’49”E, 1050 m, 13 May 1997, fl. ( MO, P, TAN); ETYMOLOGY.— The species epithet honors Randriambololona et al. 17, Mantady PN , 18°56’S, Laurent GAUTIER, our friend and colleague at the 48°26’E, 960 m, 27 Oct. 1997, fr. ( MO, P, TAN); herbarium in Geneva, who has significantly Service Forestier 10341, Mangorokely, Anosibe ,

Moramanga, [19°26’S, 48°12’E], 19 May 1954, fl. ( P). advanced our knowledge of the flora of the Prov. Toliara: Humbert 20704, Mt. Vohimavo au Nord Manongarivo Special Reserve. d’Ampasimena , bassin de la Manampanihy, [24°19’S, VERNACULAR NAMES.— Anjananjana, Zahana. 47°08’E], 830 m, 27-28 Mar. 1947, fl. ( MO, P) .

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 × 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, venation 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

axillary in the terminal leaves, 2-8-flowered, axes probably on similar substrates (Fig. 2). In the sparsely to densely puberulous; peduncle 1-2 mm past, this species has been broadly circumscribed long, distinct, involucre ellipsoid to oblong, to include all of the new taxa described here, as glabrous, 7 mm tall, 3 mm diam., apical margin well as the Sambirano endemic L. cuspidata . As with 8 narrowly triangular, subequal teeth 1 mm defined here, L. multiflora can be distinguished tall, 1 mm broad at base, glabrous or sometimes by its glabrous, elliptic, usually almost equilateral sparsely short whitish hirsute; sepals completely leaves, the largest of which usually exceed 5.5 cm included within the involucre, 4 × 4 mm, circular, in length and dry an olive green to brownish strongly inrolled, densely whitish hirsute outside, color. In addition, the sepals at anthesis exceed glabrous inside; petals narrowly oblong, 12 × the involucre teeth by (0-8-) 1-1.8 mm, a feature 3 mm, clear yellow with evident venation upon that separates L. multiflora from L. abrahamii . drying, glabrous, exserted 5-6 mm beyond involucre; stamens 6-8, free to the base, filament 8- V E R N AC U L A R N A M E S.— Amaninombi- 10.5 mm long, anthers unknown; ovary ellipsoid, lahimena, Amaninombilahy, Amaninombilahy à 1 mm diam., densely whitish golden hirsute- petites feuilles, Anjananjana, Birana, Dilatra, strigose, the trichomes to 2 mm long, style Dilatry Lahy, Fotona, Fotona Beravina, Fotonala, 11.5 mm long, slender, exserted 4-5 mm above Hazoambo, Hazomasy, Helana, Jana, involucre, stigma capitate, 1 mm in diam. Fruit Manilombilahy, Manina ombilahy, Sarifatra, unknown.— Fig. 7 . Tsilaitra à petites feuilles. CONSERVATION STATUS.— Despite an extent

Leptolaena masoalensis is known only from the of occurrence and area of occupancy far in excess type collection made at Ambato, a hill in the of 20,000 km 2 and 2,000 km 2 respectively, with northeastern part of the Masoala Peninsula (Fig. 6) only four of c. 19 sub-populations occurring in well known for its large population of Nepenthes protected areas, and thus indicating a possible masoalensis Schmid-Hollinger. At first glance, population reduction in the next three genera- L. masoalensis may be mistaken for L. delphinensis , tions of> 50%, L. multiflora is assigned a prelimwith which it shares leaves with an obtuse to inary status of Endangered (EN). rounded apex. However, L. masoalensis can be easily distinguished by its much larger, ellipsoid MATERIAL EXAMINED. — Baron 2632, Bojer s.n., to oblong involucre 7 mm tall, sepals that are Forbes s.n., Louvel 6, Thouars s.n., without precise completely included within the involucre at locality; Andrianjafy 66, Rantabe; Bernard 372, anthesis, and leaves with only 8 secondary veins Masoala PN; Birkinshaw 354, Mananara RB; Boivin s.n., Foulpointe; d’Alleizette 268, Nosy-Be(?); Decary

per side versus (8-) 11-12 in L. delphinensis . 5198, Ifandana; Humblot 349, Nosive; Perrier de la Bâthie 14186, Mahanoro; Rabe 157, Beankoraka;

CONSERVATION STATUS.— With an area of oc- Rabenantoandro 563, Mahatsara STF; Rabevohitra cupancy <100 km 2, and only a single known pop- 3808, Mahabo; Rahajasoa 424, Masoala PN; ulation that appears to lie just outside the limit of Ilaka-Est Raholivelo, 37 458, Tampolo, Tampolo STF STF; A;. Réserves Randrianasolo Naturelles 292,

Masoala National Park, L. masoalensis is assigned a 2746, 3403, 5514, 9113, 9124, 9310, Masoala RNI; preliminary status of Critically Endangered (CR). 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,

6.

Leptolaena multiflora Thouars Manampano Ampasinambo ;; Service Service Forestier Forestier 745-R- 174 1, - Ambila- R-116, Lemaitso STF; Service Forestier 1371, Andatsakala;

Hist. Vég. Isles Austral. Afriq.: 41, t. 11 (1805). — Service Forestier 1616, Ambila-Lemaitso STF; Service Type: Thouars s.n., Madagascar, without precise local- Forestier 2414, Soanierano Ivongo; Service Forestier ity ( P!). 3680, Tohakandra; Service Forestier 3809, Misevo; Service Forestier 4679, Antetezana STF; Service

As here recognized, Leptolaena multiflora Forestier 4813, Ifanirea; Service Forestier 5107, Berano; occurs primarily in coastal forest mostly on sand, Service Forestier 5677, Foulpointe; Service Forestier with a few populations recorded farther inland, 6252, Rantabe; Service Forestier 7494, Fampotakely;

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.

Figures

Fig. 4. — Leptolaena delphinensis: A, flowering branch; B, flower (McPherson et al. 14222).

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, Gautier 2361; B-E, Schatz 3233).

Fig. 7. — Leptolaena masoalensis: A, flowering branch;B, leaf (adaxial surface);C, leaf (abaxial surface);D, flower (Rahajasoa et al. 790).