Xylopia beananensis Cavaco & Keraudren, 1956

Johnson, David M. & Murray, Nancy A., 2020, A revision of Xylopia L. (Annonaceae): the species of Madagascar and the Mascarene islands, Adansonia 42 (1), pp. 1-88 : 20-22

2. Xylopia beananensis Cavaco & Keraudren ( Figs 1G ; 8J-P )

Bulletin de la Société botanique de France 103: 275 (1956). — Type: Madagascar. Prov. Toamasina [“ Domaine de l’Est ”], bassin

de la Rantabe, environs de Beanana, 650 m, 25.II.1954 (fl., fr.), Capuron 9077 (holo-, P[ P030365 ]!; iso-, P[P030366, P030367, P00697844]!; TEF, photo!; WAG!).

MATERIAL EXAMINED. — Madagascar. Prov. Antananarivo. Centre, forêt d’Ambotantely, sur le Tampoketsa d’Ankazobe, 1600 m, 27.II.1966 (fl.), Capuron 24569 SF (K n.v., P[P01954040, P01954041, P01954042]); 3.0 km NW of Ambohitsaratelo-Bebao ( NW of Tsiroanomandidy ), 1100-1200 m, 16.I.1985 (fr.), Dorr et al. 3588 ( BR, G, MO, NY, P[P01954044, P01954045], US, WAG) .

Prov. Antsiranana. Réserve naturelle intégrale de Tsaratanana, slopes W of River Antsahamanara, Fir. Marolana, Fiv. Ambanja, 14°02’06”S, 48°47’08”E, 1408 m, 28.XI.2000 (buds), Antilahimena et al. 667 ( OWU, P[P02034675]); Diana Region , Anketrabe-Belinta , Ambilobe , Beramanja, Camp Bilahy, 13°35’09”S, 48°43’34”E, 963 m, 22.II.2005 (fl.), Buerki et al. 14 ( MO) ; Ambilobe , Beramanja, Anketrabe, forêt de Kalabenono, 13°38’31”S, 48°40’25”E, 700 m, 25.XI.2006 (fr.), Callmander et al. 598 (K n.v., P[P01954043]) ; East (Nord), environs sud de Tsaratanana, entre Nosiarina et Antsirabe-Nord (route Sambava-Vohémar), 1.IV.1967 (fl.), Capuron 27657 SF (P[P01954055, P01954056, P01954057, P01954058]) ; SW of Andapa, near SW edge of Anjanaharibé-Sud Reserve , 14°48’15”E, 49°26’45”E, 1000-1100 m, 6.VIII.1997 (fr.), McPherson 17267 (K n. v., MO, WAG) ; Sava Region , Sava , Andapa , Anjialavabe, Ankiakabe, forêt dense humide de moyenne altitude d’Antsahaberaoka, située à 11 km au NW d’Anjialavabe, 14°09’53”S, 49°22’52”E, 1002 m, 24.II.2007 (fr.), Rakotovao et al. 3717 ( MO) ; Marotolana à 7 km est de Beangona-Ambevy, 14°01’54”S, 48°47’01”E, 700-1300 m, 26.XI.2000 (fl.), Ratovoson et al. 342 (P[P01954032]) .

Prov. Fianarantsoa. Fianarantsoa, Iakora, Begogo, Bekora, forêt de Sahalava au S de village du Androizaha, 23°32’26”S, 46°32’00”E, 1260 m, 29.I.2005 (fr.), Andrianjafy et al. 800 (MO).

Prov. Mahajanga. Sous-Préfecture de Befandriana-Nord, commune rurale de Matsoandakana, Quartier de Belalona, Sud-Ouest d’Andapa, Réserve Spéciale Anjanaharibe-Sud, village d’Anjiamazava, suivant la route Nationale d’Andapa-Bealanana, piste vers Nord approchant le sommet de Bevitsika, 14°42’S, 49°27’E, 1100 m, 14.XII.1994 (fr.), Ravelonarivo & Rabesonina 569 (OWU, P[P01954033]).

Prov. Toamasina. Alaotra Mangoro, Moramanga, Ambatovola, Fanovana, forêt de Vohimana, 18°55’13”S, 48°30’49”E, s. d. (fr.), Andriatsiferana 2590 (K n.v., MO, P[P00909316]); Alaotra-Mangoro Region, Fivondronana Moramanga, Commune Andasibe , Fokontany Beravo , Ambatovy forest , 18°48’28”S, 48°18’43”E, 1100 m, 26.I.2005 (fr.), Antilahimena et al. 3247 (K n.v., MO, P[P01954034]) ; Alaotra-Mangoro Region, Fivondronana Moramanga, Commune Andasibe , Fokontany Beravo , Ambatovy forest , near Berano village , 18°50’17”S, 48°19’42”E, 1025 m, 11.II.2005 (fr.), Antilahimena et al. 3377 (K n.v., MO, P[P01954031]) ; Moramanga, Andasibe, Menlamba , near Behontsa village , Ambatovy forest , 18°51’59”S, 48°19’09”E, 1045 m, 8.III.2005 (fr.), Antilahimena et al. 3623 (K n. v., P[P01954035]) ; Toamasina, Moramanga, Commune Ambohibary, Fokontany Ampitambe , 18°51’02”S, 48°17’24”E, 1080 m, 9.II.2007 (fr.), Antilahimena & Edmond 5244 (P[P06901360]) ; au-dessus d’Andratambe , alt. 500 m, bassin inférieur de la Vohilava, affluent rive gauche de la Rantabe, III.1954 (large buds, fr.), Capuron 9159 ( OWU, P[P01757050, P01954037, P01954038]) ; Alaotra-Mangoro Region, District Moramanga, Commune Ambohibary , Fokontany Ampitambe , Ambatovy , NE de Moramanga, environ 22 km de Moramanga , 18°51’31”S, 48°19’03”E, 1121 m, 19.II.2005 (fr.), Rakotovao et al. 1232 ( MO) ; Alaotra-Mangoro Region, Analamay, Ambatovy , Fokontany Ampitambe , Firaisina Ambohibary , Fivondronana Moramanga , 18°50’52”S, 48°18’31”E, 1148 m, 10.II.2005 (fr.), Razafindraibe & Antilahimena 101 ( OWU) ; Alaotra-Mangoro Region, Torotorofotsy forest, Fokontany Ampitambe , Ambatovy , commune rurale Ambohibary , district Moramanga , 18°51’48”S, 48°20’33”E, 958 m, 13.I.2005 (fr.), Razanatsoa 27 (K) ; forêt Tsiazonomby, Dist. Moramanga, Canton Anosibe , 26.VI.1952 (old fls, fr.), Service Forestier (Stanislas) 6123 SF (P[P01954049, P01954050 (fruit), P01954051]) ; Antetezampandrana, Toby Comexfo P. K. 45 route Moramanga-Anosibe, District Moramanga , Canton Moramanga , 23.I.1969 (buds), Service Forestier 26848 SF (P[P01954039]) .

DISTRIBUTION, ECOLOGY AND CONSERVATION STATUS. — Xylopia beananensis occurs in humid lowland to lower montane forests of northern, central, and southern Madagascar at elevations from 500 to 1600 meters ( Fig. 28 ). Specimens with enlarging buds have been collected in January, March, and August, with open flowers in February, April, and November, and with floral remnants in June. Specimens with mature fruits have been collected from November to March and with immature fruits in August. We have given this species a preliminary conservation assessment of Data Deficient, but its large EOO, at 192 983 km 2 the largest for any Madagascar Xylopia species ( Table 2 ), suggests that it is not of immediate conservation concern.

LOCAL NAMES. — Hazoambo (Antilahimena et al. 337, Razafindraibe & Antilahimena 101), rotry (Ravelonarivo & Rabesonina 569).

DESCRIPTION

Tree up to 30 m tall; d.b.h. up to 42 cm.

Twigs finely appressed-pubescent, the hairs 0.1-0.2 mm long, becoming glabrate; nodes with one axillary branch.

Leaves with larger blades 6.5-10.8 cm long, 2.5-4.8 cm wide, coriaceous, discolorous, olive-green and shining adaxially, tan to rusty abaxially, broadly lanceolate, oblong-lanceolate, elliptic, or rarely obovate, apex acute to acuminate, the acumen 5-15 mm long, base broadly cuneate and short-decurrent on petiole, margins somewhat revolute, glabrous adaxially, sparsely appressed-pubescent to glabrous abaxially; midrib red toward the base adaxially, secondary veins weakly brochidodromous, 10-14 per side, diverging at 45-70° from midrib, these and higher-order veins slightly raised and forming a conspicuous reticulum on both surfaces; petiole 3.5-8 mm long, flattened, winged along margins, longitudinally wrinkled, glabrous.

Inflorescences axillary or from the axils of fallen leaves, 1-4-flowered, pedunculate or not, pubescent; peduncle 1.1-2 mm long or absent; pedicels 2-4 per peduncle or arising separately from leaf axil, often recurved, 5.3-7.6(- 11) mm long, 1.0- 1.3 mm thick; bracts 2-4, spaced evenly along pedicel, caducous, only scars seen; buds lanceolate, apex obtuse.

Sepals wide-spreading at anthesis, ¹⁄5-1/2-connate, 2.7-3 mm long, 2.8-4.7 mm wide, coriaceous, semicircular, apex rounded or occasionally apiculate, pubescent abaxially.

Petals white to dull yellow, with the inner petals tinted red at the base in vivo; outer petals slightly spreading at anthesis, 17-18 mm long, 4-5 mm wide at base, 2.5-3 mm wide at midpoint, fleshy, lanceolate, keeled only at tip adaxially, with a faint keel abaxially, apex obtuse, densely puberulent except for the glabrous base adaxially, appressed-pubescent abaxially; inner petals slightly spreading at anthesis, 13.5-16.9 mm long, 3.5-3.9 mm wide at base, c. 1.2 mm wide at midpoint, fleshy, linear, keeled on both surfaces, apex acute, base concave with undifferentiated margins, densely puberulent except for the glabrous base on both surfaces.

Stamens 150-200; fertile stamens c. 1.5 mm long, narrowly oblong, anther connective apex c. 0.2 mm long, shield-shaped, slightly overhanging anther thecae, papillate, anthers 11-12-locellate, filament 0.5-0.6 mm long; outer staminodes 1.4-1.5 mm long, clavate, apex rounded; inner staminodes c. 1.1 mm long, oblong, apex truncate; staminal cone 2.8-3.2 mm diameter, 1.3-1.4 mm high, completely concealing the ovaries, rim irregularly laciniate.

Carpels c. 15; ovaries c. 0.7 mm long, ovoid, densely rusty appressed-pubescent, stigmas connivent, c. 3.3 mm long, filiform, glabrous.

Fruit of up to 15 monocarps borne on a pedicel 12-20 mm long, 2.5-3.5 mm thick, sparsely pubescent; torus 6.5-10 mm in diameter, 4-6 mm high, depressed-globose; monocarps with bright pink-red to red exterior and red endocarp in vivo, 2.4-4.8 cm long, 0.7-1.1 cm wide, 0.7-0.8 cm thick, narrowly oblong, often slightly torulose, apex obtuse, sometimes with a beak 2-5 mm long, base sessile, obliquely wrinkled, sparsely pubescent to glabrate; pericarp 0.6-2 mm thick.

Seeds up to 5 per monocarp, in a single row, oblique to parallel to long axis, 7.1-8.3 mm long, 3.7-5.2 mm wide, 3.5-4.5 mm thick, oblong-ellipsoid, circular to broadly elliptic in cross-section, brownish black to black, smooth or slightly pitted and wrinkled, shiny, perichalazal ring inconspicuous to slightly raised; sarcotesta absent; aril white in vivo, whitish orange when dried, bilobed, lobes 1.2-3.0 mm long, 3.1-4.7 mm wide, papery to fleshy, with ruffled margins.

NOTES

Xylopia beananensis is distinguished by its coriaceous broadly lanceolate to elliptic leaves with a prominent vein reticulum on both surfaces, the well-developed staminal cone, and the strongly wrinkled and narrowly oblong monocarps with seeds lying nearly parallel to the long axis. It is most similar to X. flexuosa , but differs in the flattened petioles, shorter pedicels with caducous bracts, and shorter petals. One collection (Antilahimena et al. 3377) reports the bark to be white. Precocious growth from the axillary vegetative bud superposed above inflorescences is an odd feature of some specimens.

At the time of the floristic treatment of Cavaco & Keraudren (1958) this species was only known from the basin of the Rantabe River in what is now northern Toamasina Province, but more recent collections show that it occurs up to 1600 meters from Antsiranana Province in the north south to Fianarantsoa Province. It is one of several relatively widespread mid-elevation Xylopia species on the island, including X. lemurica and X. perrieri , which do not occur below 400 meters.

Of all species of sect. Xylopia from the region, X. beananensis bears the closest resemblance, in habit, foliage, flowers, and fruits, to X. aethiopica , the single species of the section that occurs in Africa.

Figures

FIG. 1. — Aspects of Xylopia L. vegetative and floral morphology: A, shrub habit of X. humblotiana Baill.; B, habit of X. bemarivensis Diels, showing several nodes with both collateral buds forming shoots, creating pseudo-opposite branches; C, flowers of X. lastelliana Baill., showing inner petals bent outward through the gaps between outer petals (Schatz & Modeste 2877); D, flowers of X.perrieri Diels showing conspicuous red coloration at the bases of the outer petals (Schatz & O’Connor 2964); E, spreading outer petals and erect inner petals of X. kalabenonensis D.M.Johnson, Deroin, & Callmander (Razafitsalama et al. 1041); F, flower of X. richardii Boivin ex Baill., with stigmas emerging in the center amidst the inner petals; G, inflorescences of X. beananensis Cavaco & Keraudren, showing a bud, two flowers with petals, stamens, and stigmas fallen so that the staminal cone is visible, and one open flower (Buerki 14); H, open flower of X. lamarckii Baill.; I, open flower of X. lamarckii with one outer petal and one inner petal removed to show interior of flower, including stamens and stigmas. Photos: A, M. C. Andriamahay; B, F. Rakotoarison; C, D, G. E. Schatz; E, J. Razafitsalama; F, H, I, C. Kaiser-Bunbury; G, M. W. Callmander. A, B, used under the terms of the Creative Commons license for iNaturalist; C-E, G, used under the terms of the Creative Commons license for Tropicos, the botanical information system at the Missouri Botanical Garden.

FIG. 8. — Xylopia richardii Boivin ex Baill., X. beananensis Cavaco & Keraudren, and X. flexuosa Diels. A-I, X. richardii: A, stamen, abaxial view; B, flower bud, side view; C, D, leaves; E, fruit; F, seed, view of micropylar end; G, seed, side view; H, basal half of inner petal, adaxial surface; I, outer petal, adaxial surface. J-P, X. beananensis: J, outer petal, adaxial surface; K, inner petal, adaxial surface; L, flower bud, side view; M, seed, view of micropylar end; N, seed, side view; O, leaves; P, fruit. Q-X, X. flexuosa: Q, inner petal, adaxial surface; R, outer petal, adaxial surface; S, stamen, abaxial view; T, seed, view of micropylar end; U, seed, side view; V, fruit; W, leaf; X, flowers, side view. A, C, H, I, from Friedmann 3037 (MAU); B, from Florens et al. 24082 (MAU);D, from Bernardi 15071 (NY); E-G, from Lorence R 41 (MO); J-L, from isotype specimen at TAN; M, N, P, from Ravelonarivo & Rabesonina 569 (OWU); O, from Dorr 3588 (NY); Q-S, from Perrier de la Bâthie 4973 (B); T-V, from Service des Eaux et Forêts de Madagascar 12450 SF (P); W, X, from Decary 5570 (P). Scale bars: A, S, 1 mm; B, L, X, 1 cm; C-E, O, P, V, W, 2 cm; F, G, M, N, T, U, 3 mm; H-K, Q, R, 5 mm.

FIG. 28. — Distributions of Xylopia ambanjensis Cavaco & Keraudren, X. anomala, sp. nov., X. australis, sp.nov., X.beananensis Cavaco & Keraudren, X. bemarivensis Diels,X.buxifolia Baill., X.capuronii Cavaco & Keraudren,X. carinata, sp. nov., X. danguyella Ghesq. ex Cavaco & Keraudren, X. dielsii Cavaco & Keraudren, X. fananehanensis Cavaco & Keraudren, and X. flexuosa Diels on Madagascar.