A revision of Xylopia L. (Annonaceae): the species of Madagascar and the Mascarene islands Author Johnson, David M. Author Murray, Nancy A. text Adansonia 2020 2020-02-05 42 1 1 88 journal article 21758 10.5252/adansonia2020v42a1 28f94b79-38d7-4c25-ac76-cf64809114fc 1639-4798 3877215 15. Xylopia buxifolia Baill. ( Fig. 17A-K ) Adansonia; Recueil d’Observations botaniques 4: 143 (1864). — Type : Madagascar . Without definite locality, Du Petit-Thouars s. n. (holo-, P [ P030370 ]!; iso-, P[P030371]!). MATERIAL EXAMINED. — Madagascar . Prov. Antsiranana , Vallée de la Lokoho, près de Ambalavoniho [ 14.5667°S , 49.7333°E ], 75-300 m , 9-10.I.1949 (fr.), Humbert & Cours 22831 ( OWU , P[P01757073]); sous-préfecture: Antsiranana II , Commune : Antsalaka , Fokontany : Antsalaka Morafeno , Ampanihy , à 5 km à l’Ouest du village Antsalaka Morafeno , 12°36’22”S , 49°13’33”E , 717 m , 13.XII.2004 (fl.), Rakotondrafara et al. 280 ( MO ) ; Sous-Préfecture d’Andapa , Commune Rurale d’Andapa , Forêt Domaniale de Masiaposa , 14°39’20”S , 49°42’20”E , 890 m , 10.XI.1995 (fr.), Ravelonarivo & Lowry 887 (K n.v., MO , OWU ) ; Réserve Spéciale #5, Manongarivo , 30 km SE of Ankaramy , western slopes of Antsatrotro , 14°08’S , 48°21’E , 550 m , 23-25.III.1991 (fl.), Schatz 3220 (K n.v., MO ) . Prov. Fianarantsoa . Fivondronana: Mahanoro, Firaisana: Ambodibonara, Fokontany: Ambalavontaka, 20°23’26”S , 48°32’43”E , 10 m , 20.IV.2004 (fr.), Ranaivojaona et al. 645 ( OWU , P[P01953922]); Manampona, Mananjary, 7.I.1952 (fl.), Service Forestier 5613 SF (P[P01953971]); F. Manampano, Canton Mananjary, Dist. Mananjary, au bord du canal d’Ampangalana, 10 m , 27.VIII.1963 (fr.), Service Forestier 21337 SF (P[P01953944, P02133050]). Prov. Toamasina . Ampasimbe Onibe, Andranotsara, Station Forestière Mahatwara, 17°38’16”-17°38”21”S , 49°29’02”-17°29’11”E , 5-10 m , 24.IX.2002 (fr.), Andriamihajarivo et al. 72 (MO[2 sheets]); Ampanotoamaizina , 12.XI.1966 (fr.), Boiteau 505 (K) ; à Ambila- Lemaitso , 8-10.VIII.1957 (immature fr.), Capuron 18057-SF (K) ; environs de la Baie d’Antongil, W. d’Anandrovola, 100-150 m , 11.IX.1957 (fr.), Capuron 18234 SF (K, P[P01953920]); Toamasina II, Foulpointe , Morarano , Analalava , forêt d’Analalava à 6 km Sud- Ouest de Foulpointe , 17°42’19”S , 49°27’28”E , 50 m , 13.X.2004 (fr.), Lehavana et al. 142 (K n.v., MO , P[P01953964]) ; Tamatave , Tampina , Rés. For. Chemins de Fer , 5 m , 21.XII.1938 (fl.), Lam & Meeuse 6053 (K n.v., L, WAG ) ; environs d’Ambila-Lemaitso , 14.XI.1986 (fr.), Le Thomas 112 (P[P01953984, P01953985]) ; Toamasina , Ambila-Lemaitso , E of Brickaville , Old Station Forestière , c. 5 km S of town, 18°54’S , 49°07’E , 10 m , 17.I.1999 (fr.), Lowry & Miller 5138 (K n.v., MO , P[P01953987]) ; Toamasina , District : Fénérive-Est , Canton : Ampasina , Tampolo , Station forestière de Tampolo , 17°16’52”S , 49°24’33”E , 0-10 m , 2.VII.2001 (fr.), Ludovic et al. 106 (K n.v., MO , P[P01953968]) ; Toamasina , Fivondronana : Toamasina II, Firaisana : Ampasimbe , Fokontany : Mahatsara , station forestière de Mahatsara , 17°38’13”S , 49°29’03”E , 2 m , 7.VII.2001 (fr.), Ludovic et al. 118 ( MO , NY , P[P01953979]) ; Toamasina, Toamasina Dist. , vicinity of entrance to Mahatsara Forest Reserve , 1.5 km E of National Highway 5, 9.4 km N of Mahavelona village ( Foulpointe ), 17°38’S , 49°30’E , 8 m , 24-27.II.1992 (fr.), Noyes et al. 944 (K n.v., MO , P[P01953958]) ; près de Tamatave, s. d. (fr.), Perrier de la Bâthie 14912 (P[P01953941, P01953942, P01953960]); Maroantsetra , Tampolo , Péninsule Masoala , 15°43’S , 49°57’E , 20-50 m , 5.XI.2001 (fr.), Poncy et al. 1593 (K n.v., MO , P[P00373122]) ; Toamasina , Fivondronana : Mahanoro , Firaisana : Ambodibonara , Fokontany : Ambalavontaka , forêt de Nankinana , 20°22’21”S , 48°33’15”E , 41 m , 8.XI.2003 (fl.), Rabehevitra et al. 682 ( OWU , P[P01986993]) ; Toamasina , Fiv. Fénérive Est , Fir. Ampasimanigoro , Fkt. Tampolo , forêt littorale sur sable de Tampolo , 17°17’22”S , 49°24’31”E , 10 m , 30.VI.2001 (fr.), Rabenantoandro et al. 544 ( MO , NY , P[P01986995]) ; Toamasina , Fiv. Soanierana-Ivongo , Fkt. Antanambao-Ambodimanga , forêt littorale sur sable d’Antanambao 16°47’27”S , 49°43’22”E , 5 m , 31.I.2003 (fl.), Rabenantoandro et al. 1184 (K n.v., MO , NY , P[P01953921]) ; Toamasina , Soanierana-Ivongo , Antanambao-Ambodimanga , 16°46’08”S , 49°43’53”E , 18.V.2003 (immature fr.), Rabenantoandro et al. 1511 ( MO , P[P01071823]) ; region Analanjirofo , Fénérive-Est , Station forestière Tampolo , 17°16’31”S , 49°24’13”E , 10 m , 20.VI.2009 (fr.), Rakotonasolo et al. RNF 1328 (K) ; Toamasina , Tampolo ( Masoala ), Ambanizana , Anjahana , Maroantsetra , forêt de Tampolo , 15°43’45”S , 49°57’38”E , 10 m , 16.X.2001 (fr.), Randrianaivo 680 ( MO , NY , P[P01986989]) ; Toamasina , Tampolo ( Masoala ), Ambanizana , Anjahana , Maroantsetra , forêt littorale de Tampolo , 15°43’45”S , 49°57’38”E , 10 m , 18.X.2001 (fr.), Randrianaivo 705 ( MO , NY , P[P01953957]) ; District Toamasina II, Commune Rurale Ambodiriana , Fokontany Fontsimavo , village le plus proche Marinidrano , forêt de Vohidakatra , 17°56’41”S , 49°12’34”E , 294 m , 14.XI.2016 (fl.), Rasoanindriana et al. 15 ( OWU ) ; Toamasina , Nosy Mangabe , a 520 hectare island in the Bay of Antongil , 5 km S of Maroansetra , 15°30’S , 49°46’E , 200 m , 13-17.XII.1989 (fl., fr.), Schatz & Carlson 2861 ( BR , DSM , G, K n.v., MO [2 sheets], NY , WAG ) ; Tampolo–Fénérive , 3.I.1955 (fl.), Service Forestier 12497 SF (K n.v., P[P01953948]) ; Tampolo , Ampasina , Fénérive , 21.III.1957 (fr.), Service Forestier 16891 SF (P[P01953965, P01953966]) ; Toamasina , Masoala Peninsula , S of village of Ambanizana in the Andranobe River Watershed , 15°40’24”S , 49°57’51”E , 110-260 m , 29.XII.1994 (fl.), Vasey & Behasy 232 ( MO ) ; Toamasina , Masoala Peninsula , S of village of Ambanizana in the Andranobe River Watershed , 15°40’24”S , 49°57’51”E , 110-260 m , 20.XII.1994 (fl.), Vasey & Behasy 241 ( MO ) ; Toamasina , Masoala Peninsula , S of village of Ambanizana in the Andranobe River Watershed , 15°40’24”S , 49°57’51”E , 110-260 m , 22.XII.1994 (fl.), Vasey & Velo 269 (K n.v., MO ) . FIG. 17. — Xylopia buxifolia Baill. and X. lemurica Diels. A -K , X. buxifolia ; A , leaves; B , fruit; C , seed, side view; D , seed, view of micropylar end; E , outer staminode; F , fertile stamen; G , inflorescence; H , outer petal, adaxial surface; I , inner petal, adaxial surface; J , flower with petals and most of stamens removed to show staminal cone and stigmas; K , bud, side view; L -N , X. lemurica ; L , flower bud, side view; M , open flower, side view; N , habit. A -K , from Schatz & Carlson 2861 ; A , WAG; E , F , H -K , MO,sheet 1 of 2; B -D , G , MO sheet 2 of 2; L -N , from Perrier de la Bâthie 4975 (P). Scale bars:A, B, G,K-N,1 cm;C, D,5 mm;E,F,1 mm;H,I, 3 mm;J, 2 mm. Prov. Toliara . Fort Dauphin (Tôlanaro) region, c. 115 road km N of town, c. 5 km NW of Manantenina, in Forêt d’Analalava, 24°13’S , 47°21’E , 40 m , 29.X.1989 (fr.), McPherson 14285 (K n.v., MO); Taolagnaro , Région Anosy , Commune Bevoay , Mahatalaky , Sainte Luce , 24°48’46”S , 47°09’25”E , 5 m , 9.XI.2006 (fr.), Ramison 26 ( MO [2 sheets], P[P06490489]) ; Taolagnaro , Région Anosy , Commune Bevoay , Fokontany Fenoambany , 24°29’45”S , 47°05’52”E , 382 m , 17.XI.2009 (fr.), Razakamalala et al. 4639 ( MO , OWU ) ; Mandena , Fort Dauphin , 5.XII.1952 (fl.), Service Forestier 7418 (P[P01757072, P01953863]) . DISTRIBUTION, ECOLOGY AND CONSERVATION STATUS. — Xylopia buxifolia is a common component of lowland humid forests in eastern Madagascar , often occurring in littoral forests on sand, at elevations from sea level to 300 meters ( Fig. 28 ); there are scattered specimens from higher elevations (500-900 meters) in northern and northeastern Madagascar . It is a medium-sized or occasionally emergent ( Andriamampianina 2013 ) tree; a narrow crown diameter of 5-6 m for trees 21-26 m tall was reported on the labels of Vasey & Behasy 232 and Vasey & Velo 269 . In sandy littoral forest habitats it may occur together with species of Brochoneura ( Ludovic 106 ). Andriamampianina (2013) listed as additional associates Ocotea grayi , Protium madagascariensis , Ravenala madagascariensis , Rinorea arborescens , Tambourissa purpurea , and Trilepisium madagascariense at a field site in Fianarantsoa Province . Specimens with flowers have been collected in January, March, November, and December, and those with fruits from all months of the year; Andriamampianina (2013) reports flowering from November to February and fruiting from January to April at the Fianarantsoa site. With an EOO and AOO of 149 436 km 2 and 92 km 2, respectively, X. buxifolia is given a preliminary conservation assessment of Data Deficient, but does not appear to be of immediate conservation concern. Although individual local populations may be threatened, plots at three field sites showed the presence of both mature and regenerating individuals of X. buxifolia ( Andriamampianina 2013 ) . LOCAL NAMES. — Fotsy vavo ( Ramison 26 ), hazoambo ( Ludovic et al. 118 , Noyes et al. 944 , Rabenantoandro et al. 1511 ), hazoambo mena ( Ludovic et al. 106 ), hazoambo madinidravina ( Rabenantoandro et al. 1184 ), ombavy ( Ludovic et al. 118 ), ramiavona ( Service Forestier 5613 SF ). DESCRIPTION Tree up to 26 m tall, rarely a rounded shrub up to 5 m tall; d.b.h. up to 40 cm . Twigs red-brown or orange-brown pubescent, the hairs 0.1- 0.4 mm long, eventually glabrate; nodes with one or often two axillary branches. Leaves with larger blades 2.8-5.7 cm long, 1.2-2.7 cm wide, chartaceous to subcoriaceous, slightly discolorous, purplish green to light olive-brown and often somewhat shiny adaxially, dull and tan abaxially, elliptic, obovate, oblanceoleate, or occasionally lanceolate, apex obtuse to rounded and usually retuse or emarginate, base cuneate to broadly cuneate and occasionally short-decurrent on petiole, margin flat or slightly revolute, glabrous or with a few hairs along the midrib adaxially, appressed-pubescent abaxially; midrib usually darkened adaxially, secondary veins weakly brochidodromous to camptodromous, 7-13 per side, diverging at 45-60° from midrib, these and higher-order veins indistinct to slightly raised and forming a reticulum adaxially, indistinct to slightly raised abaxially; petiole 2-4 mm long, shallowly canaliculate, longitudinally wrinkled, sparsely appressed-pubescent. Inflorescences axillary, rarely supra-axillary, commonly from axils of fallen leaves, 1-4-flowered, pedunculate, pubescent; peduncle 1.2-2 mm long; pedicels 2-4 per peduncle, 4.5- 6.8 mm long, 0.5-1.0 mm thick; bracts 2-3, spaced evenly along pedicel, caducous or the distal bract sometimes persistent, 0.5-1.3 mm long, ovate; buds lanceolate, apex acute. Sepals erect to slightly spreading at anthesis, ¹⁄5-º⁄5-connate, 1.9-2.6 mm long, 1.9-2.4 mm wide, coriaceous, broadly ovate, apex acute, sparsely appressed-pubescent abaxially. Petals pale yellow, sometimes with a red blotch at the base in vivo ; outer petals slightly spreading at anthesis (ex photographs of Schatz et al. 2861 ), 9.8-16.1 mm long, 2.3-2.8 mm wide at base, 1.2-1.8 mm wide at midpoint, fleshy, linear to linear-lanceolate, keeled only at apex adaxially, keeled abaxially, apex obtuse, puberulent except for the glabrous base adaxially, sparsely appressed-pubescent or puberulent abaxially; inner petals slightly spreading at anthesis, 8.5-14.8 mm long, 1.5-1.8 mm wide at base, 0.7-0.8 mm wide at midpoint, fleshy, linear, longitudinally keeled adaxially, with a slight longitudinal keel abaxially, apex acute, base concave into a short claw, margin undifferentiated but with a transverse bilobed thickening at the widest point, puberulent except for the glabrous claw adaxially, pubescent except for the glabrous basal lobes abaxially. Stamens 100-120; fertile stamens 0.9-1.3 mm long, narrowly oblong, anther connective apex c. 0.2 mm long, capitate or shield-shaped, overhanging anther thecae, papillate, anthers 8-11-locellate, filament 0.2-0.4 mm long; outer staminodes 0.9-1.0 mm long, broadly clavate to obtriangular, apex rounded, truncate, or slightly emarginate; inner staminodes 0.8-0.9 mm long, oblong to clavate, apex truncate to slightly emarginate; staminal cone 1.1-1.4 mm in diameter, 0.6-0.7 mm high, completely concealing the ovaries, rim irregularly laciniate. Carpels 6-8; ovaries c. 1 mm long, narrowly oblong, sericeous, stigmas connivent, 1.9-3.3 mm long, filiform, glabrous to sparsely pubescent. Fruit of up to 7 monocarps borne on a pedicel 5-13 mm long, 3.5-5 mm thick, sparsely pubescent to glabrate; torus 8-14 mm in diameter, 6.3-9 mm high, depressed-globose; monocarps with greenish red, brownish red, red, or orange exterior and red endocarp in vivo , 2.3-4.0 cm long, 1.4-2.3 cm wide, 1.0- 2.2 cm thick, oblong to ovoid, not torulose, apex rounded, obtuse, or truncate, base sessile or contracted into a stipe 2-4 mm long, 5-10 mm thick, finely verrucose, obliquely wrinkled, dull to somewhat shiny, sometimes lenticellate, occasionally longitudinally ridged, glabrate; pericarp 0.8-1.5 mm thick. Seeds up to 8 per monocarp, in two rows, perpendicular to long axis, (7-) 8.5-12.6 mm long, 5.5-6.9 mm wide, 3.5-6.5 mm thick, oblong to ellipsoid but slightly narrowed toward the micropylar end and expanded into a hard donut-shaped ring, elliptic to wedge-shaped in cross-section, light brown, smooth, dull, perichalazal ring visible as a faint ridge; sarcotesta green in vivo ; aril absent. NOTES Among Madagascar Xylopia species with small but broad leaves X. buxifolia is most similar to X. lamii and X. lemurica in the needle-like inner petals and the relatively small but broad and blunt monocarps. It shares with X. lamii the obtuse to rounded and emarginate leaf apex, and the thick fruit pericarp, but it lacks the denser and more persistent reddish-brown indument of the twigs, abaxial leaf surfaces, and inflorescences of that species. The blunt leaf apex, in addition to the finer shorter hairs of twigs and leaves, distinguish it from X. lemurica , which has a pronounced leaf acumen and longer hairs on the twigs and abaxial leaf surfaces. There is, however, ecological and morphological variability within X. buxifolia as circumscribed here, and with closer field and laboratory study it is likely to prove to be a complex of species rather than a single variable one. For example, on the collection Service Forestier 12497 SF from Tampolo-Fénérive many of the leaves are acute at the apex and lanceolate to elliptic and conspicuously appressed-pubescent abaxially, but the norm for this region is obovate leaves with rounded and emarginate apices and with fine sparse appressed hairs on the abaxial surface. In the Masoala Peninsula region the leaves, while still obtuse and notched at the apex, are often larger and more lanceolate. Several specimens from Antsiranana Province , all collected from elevations above 500 meters, have subcoriaceous to coriaceous leaves that are more lanceolate and taper toward the apex while still being notched at the tip, as well as denser pubescence on the twigs and inflorescences. The single fruiting collection among these higher elevation specimens has a distinct stipe on the glabrate and wrinkled monocarps and its seeds lack the donut-shaped ring on the base of the seed, which is a consistent feature across the lowland specimens. No locality is given on the label for the type specimen. According to Lasègue (1845) , Du Petit-Thouars spent six months in Madagascar . A number of his plant descriptions were based on plant collections from Foulpointe (Mahavelona) made in 1794-1795 (Du Petit-Thouars 1804), so it is likely that the specimen was collected from that general region.