Perrierodendron capuronii, Lowry Ii, Porter P., Haevermans, Thomas, Labat, Jean-Noël, Schatz, George E., Leroy, Jean-François & Wolf, Anne-Elizabeth, 2000

Lowry Ii, Porter P., Haevermans, Thomas, Labat, Jean-Noël, Schatz, George E., Leroy, Jean-François & Wolf, Anne-Elizabeth, 2000, Endemic families of Madagascar. V. A synoptic revision of Eremolaena, Pentachlaena and Perrierodendron (Sarcolaenaceae), Adansonia (3) 22 (1), pp. 11-31 : 21-28

2. Perrierodendron capuronii J.-F. Leroy, Lowry, Haevermans, Labat & G.E. Schatz, sp. nov.

Arbor ca. 20 m alta. Folia petiolo 0.7-1. cm longo, parce stellato-pubescente; lamina obovata, chartacea subcoriaceave, (2.5-)5-9 × (2.5-) 4-6.5 cm, utrinque indumento stellato praesertim secus nervos primarios secundariosque parce induta, apice rotundata vel leviter emarginata, margine irregulariter subundulata, basi truncata subcordatave, nervis secundariis 7- ad 9-jugatis. Inflorescentia axillaris ex floribus 1 ad 7 constans, interdum in axilla inter ramulum et axin florem solitarium gerens; axe primario 1-5 cm longo, parce stellato-strigoso; pedunculis 1-3 mm longis, indumento stellato dense obtectis; involucro sub anthesi undulato, 1.5-2. mm in diam., dense papilloso-strigoso, flores sessiles unum duosve quemque ca. 2.5 cm in diam. includente. Flos sepalis 5, imbricatis, coriaceis, integris, asymmetricis, duobus externis ca. 1.5 × 1.5 mm, tribus internis ca. 8 × 6-9 mm; petalis 5, obovatis, ca. 15 × 8 mm, ut videtur glabris sed sub lente utrinque trichomatibus papillosis dispersis indutis visis, apice rotundatis emarginatis; staminibus 45 ad 50; ovario imperfecte biloculari in quoque loculo ovula duo gerente, breviter conico, a basi usque ad styli basin dense piloso, ca. 2 × 3 mm; stylo glabro, ca. 5 mm longo, ca. 1 mm in diam. Fructus obovoideus, apicem rotundatum versus leviter bilobus, 0.8-1.3 × 0.8-1.2 × 0.7-1 cm, laevis (eporcatus); exocarpio tenui dense breviter stellatostrigoso; involucro sub fructu reflexo, maturitate 0.6-1 cm in diam., irregulariter undulato; seminibus in quoque loculo uno duobusve.

TYPUS. — Service Forestier 9279, Madagascar, Prov. Antsiranana, Amponaomby, Andapa , [14°48’S, 49°33’E], 3 Dec. 1953, fl. (holo-, P!; iso-, P!, TEF) .

Trees 20 meters tall, bole to 8 m, bark with slight cracks. Young twigs with dense stellate indumentum toward apex. Leaves obovate, brown above, greenish-brown below (in dry material), chartaceous to subcoriaceous, (2.5-)5-9 × (2.5-) 4-6.5 cm, with sparse stellate indumentum mostly on the primary and secondary veins of both surfaces, apex rounded to slightly emarginate, margin irregularly and weakly undulate, base truncate to subcordate, venation brochidodromous, with 7-9 pairs of alternate to subopposite secondary veins, midrib weakly channeled above, raised below; petiole 0.7-1 cm long, sparsely stellate, adaxially canaliculate; stipules unknown, caducous, scars inconspicuous. Inflorescences axillary, with 1-7 flowers, primary axis 1-5 cm long, sparsely stellate-strigose, sometimes also with a single axillary flower borne between the stem and the inflorescence axis; bracts not seen; peduncles 1-3 mm long, densely covered with stellate indumentum, involucre in flower undulate, 1.5-2 mm in diam., densely papillate-strigose, containing 1 or 2 sessile flowers each ca. 2.5 cm in diam.; sepals 5, imbricate, coriaceous, entire, asymmetrical, the 2 small external ones 1.5 × 1.5 mm, deltoid, the 3 internal ones 8 × 6-9 mm, glabrous and papyraceous where covered by the adjacent sepal, densely stellate-strigose below where exposed; petals 5, con- torted in opposite direction from sepals, obovate, 15 × 8 mm, appearing glabrous but with scattered papillose hairs on both surfaces, apex rounded, notched; extrastaminal disc cupuliform, 1 mm high, with an irregular margin; stamens ca. 45-50, not persistent in fruit, filaments ca. 1 cm long, glabrous, anthers 1 × 1 mm, basifixed; ovary shortly conical, densely pilose (glabrous at the base of the style), 2 × 3 mm, with 2 incomplete locules each containing 2 ovules; style glabrous, ca. 5 mm long, 1 mm in diam., stigma 1-1.5 mm long, crateriform, margin distinctly undulate. Fruits obovoid, weakly bi-lobed toward the rounded apex, 0.8-1.3 × 0.8-1.2 × 0.7-1 cm, smooth (without ridges), exocarp thin, densely short stellate-strigose, endocarp thin, involucre reflexed, 0.6-1 cm in diam. at maturity, irregularly undulate; seeds 1-2 per locule, spherical to hemispherical (depending on the number), 5-6 mm long, brown, endosperm lacking, cotyledons contorted, radicle included, ca. 5 mm long. — Fig. 5 .

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Perrierodendron capuronii is known from only three collections, the type gathering, which is in flower, and two paratypes, one with numerous fruits and only a few partially damaged, detached leaves, and another from the same area with a few very young fruits and numerous well preserved leaves that closely match those of the holotype. We associate the holotype with the other collections based primarily on similarities in their inflorescence structure and leaf color, texture and venation, as well as the fact that the blade is distinctly shiny on its adaxial surface. The sites at which the paratype collections were made near Moramanga are over 400 km to the SSW of the type locality ( Fig. 4 ), and may be situated closer to sea level. However, until additional material at other stages of development becomes available from either or both areas, it seems best to treat them as belonging to a single species.

Perrierodendron capuronii can be distinguished from other members of the genus by its leaves with a shiny adaxial surface, its obovoid and weakly bi-lobed, 2-seeded fruit covered with very short stellate indumentum, and its reflexed involucre with an undulate margin.

ETYMOLOGY. — The specific epithet honors the renowned forest botanist René CAPURON, who contributed so much to our understanding of the Malagasy flora, and who took a special interest in the taxonomy and morphology of the island’s endemic plant families.

VERNACULAR NAMES. — Ampaliala, Tanantanampotsy.

PARATYPES. — MADAGASCAR, Prov. Toamasina: SF 25523, Anosibe an’Ala, Ambatoafo, Fiadanana , [19°32’S, 48°30’E], 18 Feb. 1965, ster. (K, MO, P, TEF); SF 26658 , Moramanga , Toby-Andasimanara , [19°04’S, 48°14’E], 900 m, 6 Feb. 1968, fr. ( P [2 sheets], TEF) .

3. Perrierodendron occidentalis J.-F. Leroy, Lowry, Haevermans, Labat & G.E. Schatz, sp. nov.

Arbor 3-20 m alta. Folia petiolo (0.3-)0.5-1(-1.5) longo, indumento stellato obtecto; lamina obovata usque elliptica, chartacea, 3-6.5(-8) × 2-5 cm, utrinque indumento stellato parce induta, apice truncata retusa obtusave, margine integra vel irregulariter subundulata, basi truncata obtusa emarginatave, nervis secundariis 5- ad 10-jugatis. Inflorescentia axillaris ex floribus 1 ad 5 constans, aliquando in axilla inter ramulum et axin florem solitarium gerens; axe primario 0.5-2 cm longo, parce stellato-strigoso; pedunculis 1-5 mm longis, indumento stellato dense obtectis; involucro sub anthesi ca. 2 mm in diam., dense papilloso-strigoso, florem sessilem solitarium ca. 2.5 cm in diam. includente. Flos sepalis 5, imbricatis, coriaceis, asymmetricis, duobus externis ca. 3 × 1 mm, tribus internis 5-6 × ca. 4 mm; petalis 5, obovatis, 8-9(-10) × ca. 6 mm, glabris, apice rotundatis emarginatis, basi breviter unguiculatis; staminibus 50 ad 55 in fructu persistentibus; ovario imperfecte biloculari in quoque loculo ovula duo gerente, breviter conico, a basi usque ad styli basin dense piloso, ca. 3.5 × 3 mm; stylo glabro, ca. 3.5 mm longo, ca. 1 mm in diam. Fructus late obovoideus, apice in rostrum brevem conicum productus, 1.2-1.5 × 0.7-0.9 cm, exocarpio tenui, crustaceo, utrinque laevi (eporcato), trichomatibus stellato-strigosis ramulos 1-2 mm longos gerentibus vestito; involucro maturitate ca. 0.7 cm in diam. cum staminium reliquiis persistentibus cupulam sub fructu formante; seminibus in quoque fructu uno duobusve.

TYPUS. — Service Forestier (Capuron) 20603, Madagascar, Prov. Toliara, forêt d’Analamarina (Hazoroa) au sud-est de Sakaraha , [23°00’S, 44°33’E], 500-600 m, 28 Dec. 1961, fr. (holo-, P!; iso-, P!, TEF) .

Trees 3-20 m tall. Young twigs with stellate indumentum. Leaves obovate to elliptical, dull brown above, dull green below (in dry material), chartaceous, 3-6.5(-8) × 2-5 cm, with scattered stellate indumentum on both surfaces, apex truncate or retuse to obtuse, margin subentire, slightly undulate, base truncate to obtuse or emarginate, venation brochidodromous, with ca. 5-10 pairs of alternate to subopposite secondary veins, midrib weakly channeled above, raised below; petiole (0.3-)0.5- 1(-1.5) long, adaxially canaliculate, covered with stellate indumentum; stipules linear, caducous, 1.5- 2 mm long. Inflorescences axillary, with 1-5 flowers, primary axis 0.5-2 cm long, sparsely stellate-strigose, occasionally with an axillary flower borne between the stem and the inflorescence axis; peduncles 1-5 mm long, densely covered with stellate indumentum; bracts minute (?), caducous; involucre in flower ca. 2 mm in diam., densely papillate-strigose, containing a single sessile flower ca. 2.5 cm in diam.; sepals 5, imbricate, coriaceous, asymmetrical, the 2 small external ones 3 × 1 mm, narrowly triangular, the 3 larger internal ones ca. 5- 6 × 4 mm, glabrous and papyraceous where covered by the adjacent sepals, densely stellate strigose below where exposed;petals5,contorted in opposite direction from sepals, obovate, ca. 8-9(-10) × 6 mm, glabrous, apex rounded, notched, claw short; disc cupuliform, 0.5 mm high, margin entire; stamens ca. 50-55, persistent in fruit, filament 5 mm long, anthers 1 × 1 mm, basifixed; ovary shortly conical, densely pilose (glabrous at the base of the style), 3.5 × 3 mm, with 2 incomplete locules each containing 2 ovules; style glabrous, 3.5 mm long, 1 mm in diam., stigma 1 mm long, crateriform, margin distinctly undulate. Fruits broadly obovoid, 1.2-1.5 × 0.7-0.9 cm, apex extended into a short conical rostrum, exocarp thin, crustaceous, smooth on outer and inner surfaces, without ridges, branches of stellate-strigose indumentum 1-2 mm long, transparent yellow with a reddish core, endocarp thin, crustaceous; involucre forming a cup with persistent remains of the stamens, 0.7 cm in diam. at maturity; seeds 1-2 per fruit, spherical or hemispherical, 6-9mm in diam.,brown, cotyledons very contorted, radicle included, 3-4 mm long. — Fig. 6 .

Perrierodendron occidentalis is widely distributed in western Madagascar, extending from Isalo PN to Ambalabe and Analamary in the S, with an outlying population at Manongarivo SSE of Soalala in the NW ( Fig. 4 ). It occurs on sandy substrates in seasonally dry, deciduous forests. With the exception of one collection made in 1973 and another last year, P. occidentalis has not been recorded since the early 1960s. This species closely resembles P. boinense , but can be distinguished from it by having smooth (unribbed) fruit with the apex extended into a short, conic rostrum, a thin, crustaceous endocarp, and sepals ca. 5-6 mm long and petals ca. 8-9(-10) mm long.

VERNACULAR NAMES. — Fotona, Nato, Tsiandala, Tsitaka.

PARATYPES. — MADAGASCAR, Prov. Fianarantsoa: Cremers 3584, Ambasy , ouest Ambalavao , [21°28’S, 46°12’E], 900-1000 m, 9 Dec. 1974, fl. ( P [2 sheets], TAN); Decary 16384, Isalo PN, [22°08’- 22°40’S, 45°10’- 45°24’E], 2 Nov. 1940, fl. ( P) ; Humbert 19511, Isalo PN, sud de Sahanafa, [22°20’S, 45°18’E], 1000 m, 28 Nov.-4 Dec. 1946, fl. ( P [2 sheets]); SF 4- R-258, Lalanandro, [22°18’S, 46°06’E], 29 Jan. 1925, ster. ( P); SF 59 -R-141, Ambondrona, [21°54’S, 46°52’E], 9 June 1954, ster. ( P); SF 74 -R-47, Lalanandro, [22°18’S, 46°06’E], 1 June 1952, ster. ( P); SF 239 -R-239, Isalo PN, [22°08’- 22°40’S, 45°10’- 45°24’E], 23 July 1955, ster. ( P); SF 13761 , Massif d’Ingaro, Ambalavao, [21°36’S, 46°22’E], 13 Feb. 1955, fr. ( P, TEF), SF 13878 , Isalo PN, [22°11’- 22°43’S, 45°10’- 45°21’E], 22 Jul. 1955, ster. ( P, TEF) . Prov. Mahajanga: Perrier de la Bâthie 1644, Manongarivo (Ambongo), [16°16’S, 45°22’E], Nov. 1903, fl. ( P) . Prov. Toliara: Humbert 19395, bassin de la Malio (affluent de Mangoky), près d’Ambalabe, [21°58’S, 45°15’E], 400-450 m, 23-27 Nov. 1946, fl. ( P [2 sheets]); Randrianaivo et al. 315, Zombitse PN, 12 km E of Sakaraha, 1 km N of Andranomaitso, 23°53’07”S, 44°38’42”E, 600-800m, 4 Feb. 1999, fr. (G, K, MO, P, TAN); SF 5 -R-350, Sakaraha STF, [22°53’S, 44°32’E], 24 Feb. 1956, ster. ( P) ; SF (Capuron) 20599, Analamary, forêt d’Analamarina ( Hazoroa ), au sud-est de Sakaraha , [23°00’S, 44°33’E], 28 Dec. 1961, bud ( P) .

4. Perrierodendron quartzitorum J.-F. Leroy, Lowry, Haevermans, Labat & G.E. Schatz, sp. nov.

Frutex arbusculave 3-6 m alta. Folia petiolo 0.2-0.5 cm longo, indumento fusco stellato-strigoso dense obtecto; lamina obovata usque elliptica, chartacea subcoriaceave, 2.5-4(-5) × 2-3.5(-4) cm, utrinque indumento fusco stellato dense induta, apice rotundata vel leviter emarginata, mucronata, margine integra, basi truncata usque obtusa, nervis secundariis 7- ad 10- (ad 13-) jugatis. Inflorescentia axillaris, contracta, plerumque ex floribus 1 ad 3 constans; axe primario 3-6 cm longo, dense stellatostrigoso; pedunculis longitudine 1 mm non attingentibus, dense papilloso-strigosis; involucro sub anthesi perparvo vel absente, si adest flores sessiles quemque ca. 3.5 cm in diam. includente. Flos sepalis 5, imbricatis, coriaceis, integris, asymmetricis, duobus externis 2-3 × ca. 1 mm, tribus internis 10-11 × 5-6 mm, unoquoque in partibus sepalo vicino tectis glabro papyraceoque, in partibus expositis dense stellato-strigoso; petalis 5, obovatis, 15-17 × ca. 14 mm, glabris, apice rotundatis emarginatis, mucronatis; staminibus ca. 20, caducis; ovario imperfecte biloculari in quoque loculo ovula duo gerente, conico, a basi usque ad styli basin dense piloso, ca. 3 × 3 mm; stylo glabro, ca. 2 mm longo, ca. 1 mm in diam. Fructus conicus, apice acutus, 1.9-2.5 × 0.9-1.1 cm, indumento papilloso stellato dense obtecto; endocarpio crustaceo; involucro in fructu parvo vel absente; semine in quoque fructu uno (vel rare seminibus duobus).

TYPUS. — Labat, Haevermans, Rabenantoandro & Cook 3035, Madagascar, Prov. Fianarantsoa, Ambatofinandrahana, Itremo, forêt claire sclerophylle à Perrierodendron et Leptolaena sur sol sableux sur schiste, 20°35’31”S, 46°35’43”E, 1560 m, 26 Mar. 1999, fl., fr. (holo-, P!; iso-, G!, K!, MO!, P [2 sheets]!, TAN!).

Shrubs to small trees 3- 6 m. Twigs with stellate indumentum, internodes 1.5-3(-4) mm long, giving the stems an evident “zigzag” appearance. Leaves elliptical to obovate, dark green above, pale green below (in dry material), chartaceous to subcoriaceous, 2.5-4(-5) × 2-3.5(-4) cm, with dense stellate brown indumentum on both surfaces, apex rounded to slightly emarginate, mucronate, margin entire, base truncate to obtuse, venation eucamptodromous, with 7-10 (-13) pairs of alternate to subopposite secondary veins, midrib and secondary veins weakly chan- neled above, raised below; petiole 0.2-0.5 cm long, densely brown stellate-strigose, adaxially canaliculate; stipules not seen. Inflorescences axillary, contracted, generally with 1-3 flowers, primary axis 3-6 cm long, densely stellate-strigose; peduncles very short, less than 1 mm long, densely papillate strigose; involucre in flower very small to absent, containing 2 sessile flowers each ca. 3.5 cm in diam.; sepals 5, imbricate, coriaceous, entire, asymmetrical, the 2 small external ones, 2-3 × 1 mm, narrowly triangular, the 3 internal ones 10-11 × 5-6 mm, glabrous and papyraceous where covered by the adjacent sepal, densely stellate strigose where exposed; petals 5, contorted in opposite direction from sepals, obovate, 15-17 × 14 mm, glabrous, apex rounded, notched, mucronate, claw short; extrastaminal disc cupuliform, 1 mm high; stamens ca. 20, not persistent in fruit, filaments 4-5 mm long, anthers 1 × 1 mm, basifixed; ovary conical, densely pilose (glabrous from the base of the style), 3 × 3 mm, with two incomplete locules each containing 2 ovules; style ca. 2 mm long, 1 mm in diam., forming an extension of the conical ovary, stigma 1 mm long, crateriform, margin distinctly undulate. Fruits conical, apex acute, 1.9-2.5 × 0.9-1.1 cm, with dense papillate stellate indumentum; endocarp crustaceous, involucre small to absent; seeds 1(-2), spherical to hemispherical, ca. 0.8 cm in diam., light brown (in spirit material), aborted ovules 3, persistent, embryos 1-2 per seed, cotyledons very contorted, radicle included, 1 mm long. — Figs. 7 , 8 .

Perrierodendron quartzitorum is restricted to the Itremo massif ( Fig. 4 ), where it has been collected several times recently. This species is easily recognized by the dense, stellate indumentum that covers the abaxial surface of its leaves, its short internodes that give an evident “zig-zag” appearance to the stems, and its fruit whose apex tapers into a prominent rostrum about half the length of the fruit body.

VERNACULAR NAMES. — Kitoto, Kitotovavy.

PARATYPES. — MADAGASCAR, Prov. Fianarantsoa: Du Puy, Labat & Andriantiana M 636, W of Ambositra, Itremo , 20°35’28”S, 46°35’40”E, 1500 m, 21 Nov. 1993, fl. (K, MO, P, TAN, TEF); Humbert

30023, montagnes à l’Ouest d’Itremo, [20°28’- 20°49’S, 46°26’- 46°41’E], 1500-1700 m, 17-22 Jan. & 18-22 Apr. 1955, fl. ( P) ; Service Forestier (Capuron) 11567, Monts Ambatomenaloha, ouest d’Itremo, audessus d’Antanimena, [20°32’S, 46°33’E], 20 Jan. 1955, y. fr. (K, MO, P [2 sheets], TEF), SF (Capuron) 28902, environs du col de l’Itremo, ouest d’Ambatofinandrahana, [20°35’S, 46°35’E], 1600- 1700 m, 27 Nov. 1969, fl. ( P, TEF) , SF (Capuron) 29045, montée du col de l’Itremo, Ambatofinandrahana, rive droite de l’Ambatorangotana, [20°35’S, 46°35’E], 1650-1700 m, 21 Feb. 1970, fr. (B, BR, G, K, MO, P [2 sheets], WAG) .

Figures

Fig. 5. — Perrierodendron capuronii: A, flowering branch; B, flower at anthesis; C, flower (schematic transverse section); D, sepal; E, petal; F, fruit. (A-E, Service Forestier 9279; F, Service Forestier 26658).

Fig. 4. — Distributions of Perrierodendron, mapped on the simplified bioclimatic zones of Madagascar (SCHATZ in press, after CORNET 1974). P. boinense (*), P. capuronii (+), P. occidentalis (•), P. quartzitorum (Δ), P. rodoense (-).

Fig. 6. — Perrierodendron occidentalis: A, flowering branch; B, leaf (abaxial surface) and flower buds; C, flower at anthesis; D, flow- er (some parts removed to show gynoecium); E, petal; F, fruits. (A-E, Perrier de la Bâthie 1644; F, Service Forestier 20603).

Fig. 7. — Perrierodendron quartzitorum: A, flowering branch; B, young leaf and buds; C, flower at anthesis; D, petal; E, androecium and gynoecium; F, gynoecium (portion of wall removed to show ovules); G, fruiting branch. (A-G, Labat et al. 3035).

Fig. 8. — Photographs of Perrierodendron quartzitorum: A, habit; B, mature fruit; C, flowering branch; D, flower. (A, B, D, Labat et al. 3035; C, Du Puy et al. M636). — Photos J.-N. LABAT.