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 14. Xylopia bemarivensis Diels ( Figs 1B ; 2D ; 14A-K ; 16B ) Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 9: 352 (1925). — Type : Madagascar . Prov. Mahajanga [“Madagascaria occidentalis”], Boina : ad rivum Bemarivo superiorem, III.1907 (fr.), Perrier de la Bâthie 4945 (holo-, B!; iso-, P[P00524385]!). Xylopia decidua Diels, Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 9: 353 (1925). — Type : Madagascar . Prov. Mahajanga [ “Madagascaria occidentalis ”], Boina , bois sablonneux, environs du Mt. Tsitondraina [ Tsitondroina ], XII.1900 (fl., fr.), Perrier de la Bâthie 1167 (holo-, B !; iso-, P[P030368, P030369]!). MATERIAL EXAMINED. — Madagascar . Prov. Antsiranana , Réserve spéciale d’Ankarana, à 10 km à l’Est du village de Matsaborimanga (Campement des Anglais), 12°54’59”S , 49°06’24”E , 180 m , 9.VI.1995 (fr.), Andrianantoanina et al. 848 ( OWU ) ; Ambilobe , Mahamasina , Réserve spéciale d’Ankarana , 12°57’49”S , 49°07’28”E , 100 m , 5.XI.1997 (fr.), Bardot-Vaucoulon 915 (K n.v.; MO ; P[P00309729, P00309730]) ; Ambilobe , Ambondromihefy , Réserve spéciale d’Ankarana , Mahoro , sentier botanique partie nord est en direction d’Ambalihatsara , 12°51’21”S , 49°13’45”E , 300 m , 22.I.2003 (fl.), Bardot-Vaucoulon et al. 1263 ( MO , P[P00455560]) ; Vovo Village District , c. 6 km from Diego Suarez on Diego Suarez-Vovo village road, and c. 1.5 km S of beach on Indian Ocean , 12°19’05”S , 49°23’07”E , 85 m , 21.IV.1993 (fl., fr.), Harder et al. 1658 ( MO , WAG ) ; Diana Region , Ambilobe , Ambakirano , Behefaka , Anjahana , 13°21’56”S , 49°10’04”E , 151 m , 5.VI.2005 (fr.), Hong-Wa 303 ( MO , P[P01953785]) ; Ampasindava , forêt d’Andranomatavy , 13°39’59”S , 47°59’24”E , 285 m , 25.XI.2009 (bud, fr.), Madiomanana et al. MAD 267 (K n.v., MO , P[P0175075]) ; Sava Region , Sous-préfecture Vohemar , Commune Nosibe , Fokontany Anjiabe , Village Anaborano , 13°04’06”S , 49°54’27”E , 49 m , 7.XII.2004 (fr.), Manjakahery & Sola 72 ( MO ) ; 40 km N of Vohemar , near N end of Lac Sahaka , Fir. : Nosibe , Fkt. : Anjiabe , 13°04’45”S , 49°54’17”E , 20 m , 3.XI.2002 (buds), McPherson 18833 ( OWU ) ; Analamazava , part of Binara Range , SW of Daraina ( Vohemar ), 13°15’S , 49°38’E , 200-1180 m , 26.IV.2000 (fr.), Meyers 92 ( OWU ; P[P02005891]) ; sous-préfecture de Vohemar , commune rurale de Daraina , Daraina , forêt d’Antsaharaingy , 12°55’N , 49°40’E , 50 m , 28.II.2005 (fl.), Nusbaumer & Ranirison LN 1509 (G, K n.v., MO ) ; Fivondronana : Vohémar , Firaisana : Tsarabaria , Fokontany : Manakana , 13°43’06”S , 50°05’52”E , 15.VII.2003 (fr.), Rabehevitra et al. 358 ( MO , OWU , P[P02133052]) ; Fivondronana : Vohémar , Firaisana : Nosy be, Fokontany : Anjiabe , 13°04’43”S , 49°54’04”E , 11.V.2004 (fr.), Rabehevitra et al. 938 ( OWU , P[P01954139]) ; Fivondronana : Vohémar , Firaisana : Tsarabaria , Fokontany : Manakana , 13°42’46”S , 50°05’25”E , 3 m , 18.V.2004 (fr.), Rabehevitra et al. 1009 ( MO , OWU , P[P01954128]) ; Fiv. : Vohémar , Fir. : Tsarabaria , Fok. : Manakana , 13°42’58”S , 50°05’37”E , 3 m , 24.X.2002 (fl.), Rabenantoandro et al. 1035 ( MO , OWU ) ; Fivondronana : Vohémar , Firaisana : Nosibe , Fokontany : Anjiabe , 13°04’43”S , 49°54’04”E , 10 m , 23.II.2003 (fl., fr.), Rabenantoandro et al. 1318 ( MO , OWU , P[P06901368]) ; Fivondronana : Vohémar , Firaisana : Tsarabaria , Fokontany : Manakana , 13°42’49”S , 50°05’13”E , 5 m , 1.III.2003 (bud, fr.), Rabenantoandro et al. 1380 ( MO , K n.v., OWU , P[P01987003]) ; Diana Region , Diana , Ambolobozobe , Ambovomavo , à 1 km W du village d’Ambolobozobe , 12°31’13”S , 49°31’21”E , 24.XI.2007 (fr.), Rakotonandrasana 1217 ( OWU , P[P02090377]) ; Vohemar , Maromokotra , Antsiranana , forêt d’Andavaposa , à 5 km au Sud du village d’Antsaharaingy , 12°54’05”S , 49°38’45”E , 36 m , 11.XI.2005 (fr.), Rakotondrafara et al. 430 ( MO ) ; Montagne d’Ambre , versants ouest, 12°35’12”S , 49°05’52”E , 671 m , 25.I.2012 (fl., fr.), Ramandimbimanana & Randimbiarison SDR 318 (K n.v., MO ) ; Diana Region , Antsiranana II , Ankarongana , 12°36’18”S , 49°26’34”E , 258 m , 7.XI.2006 (fr.), Ranaivojaona et al. 1527 ( OWU , P[P02090430, P02090431]) ; Diana Region , Antsiranana II, Ankarongana , Sahafary , 12°36’18”S , 49°26’34”E , 258 m , 4.XII.2006 (fl., fr.), Ranaivojaona et al. 1651 ( MO ) ; Fivondronana : Diego II , Commune : Mahavanona , Montagne des francais, forêt d’Ampitiliantsambo , à 3 heures de marche à pied, à l’Est d’Andranomanitra , 12°23’13”S , 49°23’04”E , 205 m , 14.I.2005 (fl., fr.), Randrianaivo et al. 1155 (K n.v., MO , P[P02133047]) ; Sava Region , Sous-préfecture Vohemar , Commune Nosibe , Fokontany Anjiabe , Village Anabovano , Campement Andrainginalo , 10 km à l’Est d’Anaborano , 13°04’09”S , 49°54’08”E , 18 m , 11.X.2004 (bud, fr.), Randrianarivelo et al. 159 (K n.v., MO ) ; Diana Region , Sous préfecture Antsiranana II, Commune Mahavanona , Fokontany Andranomanitra , 12°22’05”S , 49°19’37”E , 382 m , 8.II.2005 (fl.), Randrianarivelo et al. 219 (K n.v., MO ) ; Réserve Spéciale d’Ankarana , sur le Tsingy près de la grotte de chauve-souris et aux alentours, 12°54’43”S , 49°06’39”E , 180 m , 20.II.1994 (fl., fr.), Randrianambolona et al. 48 ( MO ) ; sous-préfecture de Vohemar , commune rurale de Daraina , Daraina , forêt d’Antsahabe , 13°13’N , 49°33’E , 345 m , 4.V.2004 (fr.), Ranirison PR 808 (G, K n.v., MO ) ; sous-préfecture de Vohemar , commune rurale de Daraina , Daraina , forêt d’Ankaramy , 13°18’N , 49°40’E , 24.II.2004 (fl.), Ranirison & Nusbaumer PR 475 (G, K n.v., MO ) ; Andrafiabe , Ambolobozobe , 1 km au Nord est d’Ambolobozobe , forêt d’Ampanasagna , 12°31’01”S , 49°32’14”E , 50 m , 1.II.2005 (fl.), Ratovoson et al. 864 (K n.v., MO ) ; Vohemar , Maromokotra , Ankijomantsina , 3 km au Sud d’Ankijomantsina, 13°07’40”S , 49°27’53”E , 100 m , 30.X.2005 (fr.), Ratovoson et al. 1045 ( MO , P[P01953997]) ; Diana Region , Sadjoavato , forêt de Sahafary , 12°36’26”S , 49°26’43”E , 280 m , 8.I.2007 (fl.), Ratovoson et al. 1180 ( MO , OWU , P[P01954133]) ; Ambobaka , 300 m Sud du pont Antomboka , Commune Ambodimanga Ramena , Fivondronana Ambanja , 13°45’12”S , 48°31’19”E , 41 m , 4.XII.2000 (fr.), Razakamalala et al. 49 ( MO ) ; Diana Region , Commune Sadjoavato , Fok. Saharenana , forêt d’Andranomadiro SW of Sahafary , 12°36’18”E , 49°26’35”E , 300 m , 14.II.2005 (fr.), Schatz et al. 4271 ( MO ) ; Benavone , Ambanja, Service Forestier 10275 ( OWU ) . Prov. Mahajanga . Maevatanana, Antsiafabositra, Bemanevika, 17°13’28”E , 46°59’37”E , 350 m , 7.V.2005 (fr.), Andrianjafy et al. 1003 ( MO , P[P01953993]); Beanka , partie centrale, Antsinginarefy , 17°57’23”S , 44°28’06”E , 329 m , 11.II.2012 (fr.), Bolliger et al. RFB 212 ( MO ) ; forêt d’Amparihymikimbo , auprès de l’embouchure de la Mafaika ( E de la Baie de Narinda ), 28.V.1958 (fr.), Capuron 18540-SF (K) ; forêt d’Ambondro-Ampasy ( Exploitation Loyseau ), Canton d’Antonibe , District d’Analalava , 29.X-3.XII.1958 (fl.), Capuron 18798-SF (K) ; Ambongo , sables, près de Benetsy (riv. g. de la basse Mahevavy ), 21.XI.1965 (fl.), Capuron 24263-SF (K) ; Station forestière d’Ampijoroa , c. 3 km N d’Andranofasika , 16°20’S , 46°51’E , 12.IV.1984 (fr.), Dorr & Koenders 3012 ( GH , K n.v., MO , WAG ) ; région Melaky , Beanka , partie sud, 18°01’59”S , 44°28’33”E , 230 m , 7.XII.2012 (fr.), Gautier et al. LG 5964 (K n.v., MO ) ; forêt de Tsiampihy , de Besaraha , de Bemiha et de Soahanina , Leandri et al. 2229 ( OWU ) ; Melaky Region , Maintirano , Belitsaka , Ambinda , 18°02’38”S , 44°28’56”E , 232 m , 20.X.2009 (fl.), Letsara et al. LRK 898 ( CAS , MO , P[P00722348]) ; Boriziny ( Port-Berg. ), Bongolava , on Bongolava mountain , 3 km bifurcation from Port-Bergé , 15°36’49”S , 47°35’21”E , 215 m , 18.III.2010 (fr.), Rajaovelona & Rakotonasolo RJL 166 (K) ; Station Forestière d’Ampijoroa , piste vers Jardin botanique A[mpijoroa], 16°18’40”S , 46°48’00”E , 200 m , 9.III.1996 (fr.), Rakotomalaza et al. 636 (K n.v., MO , NY ) ; Sofia Region , Boriziny , Port-Bergé , Tsingingia , Marosely , 18 km au Sud Boriziny, 15°38’58”S , 47°35’03”E , 217 m , 16.XI.2004 (fl., fr.), Ramananjanahary 121 (K n.v., MO ) ; Sofia Region , Boriziny , Port-Bergé , Tsingingia , Andranomena II , 15°34’00”S , 47°29’55”E , 115 m , 27.XI.2004 (fl., fr.), Ramananjanahary 151 ( MO , P[P01954134]) ; Faritany Mahajanga , Fivondronana Port-Bergé , Bongolava , 17°38’58”S , 47°35’03”E , 217 m , 17.XI.2004 (buds, immature fr.), Razakamalala et al. 1743 ( MO , P[P00903552]) ; Faritany : Mahajanga , Fivondronana : Port-Bergé , Mampikony , Bongolava , Betaramahamay , forêt sèche sur sable d’Ambohimanga , 15°57’05”S , 47°26’09”E , 232 m , 6.XII.2004 (fl., fr.), Razakamalala et al. 1834 (A, MO ) ; Bevazaha , canton Tsaramandroso , district Ambato Boeni , 28.XI.1951 (fl.), Réserves Naturelles 3338 (K as “ 338 ”, MO , OWU ) ; Ampijoroa , Marovoay , 16.XII.1953 (fl.), Service Forestier 8066 (K n.v., OWU ) ; forêt Katsijy , Kandreho , Maevatanana , 12.XI.1953 (fr.), Service Forestier 8073 ( OWU ) ; Ampijoroa , Marovoay , 17.XII.1953 (fl., fr.), Service Forestier 8103 ( OWU ) ; forêt Mafaijijo , Maintirano , 28.II.1956 (fl.), Service Forestier 12646 ( OWU ) ; Andrianonakanga , Sitampihy , Ambato Boini , 26.IV.1955 (fr.), Service Forestier 14921 ( OWU ) ; Jardin botanique d’Ampijoroa , Ampijoroa , Fokontany Andranofasika , Fivondronana Marovoay , 16°18’S , 46°49’E , 200 m , 25.I.1992 (fr.), Service Forestier ( Comtet ) SF 34395 (K n.v., MO ) . Prov. Toliara . Atsimo-Andrefana Region, Beroroha à 4 km avant Antsoa, 21°15’43”S , 45°10’04”E , 461 m , 3.XII.2010 (buds, immature fr.), Andriantiana et al. 1011 (MO) ; Atsimo-Andrefana Region , Makay Massif , Lake Anosilahy , 21°20’31”S , 45°10’57”E , 415 m , 27.XI.2010 (fl., fr.), Phillipson et al. 6251 ( MO ) ; Atsimo-Andrefana Region , forêt d’Anosilamy , Fokontany Beronono , Commune Beronono , 21°20’30”S , 45°10’53”E , 448 m , 13.I.2010 (fr.), Razakamalala et al. 5146 ( MO , P[P02090409]) . DISTRIBUTION, ECOLOGY AND CONSERVATION STATUS. — Xylopia bemarivensis occurs in northern and western Madagascar , in dry deciduous forest, semi-deciduous forest, and rarely littoral forest on brown or white sand, rarely on clay soils, at elevations from sea level to nearly 700 meters ( Fig. 28 ). Its distribution is discontinuous ( Fig. 28 ), reflecting an actual discontinuity in this vegetation type ( Moat & Smith 2007 ); specimens from the two areas are very similar. Occurring south of 21°S latitude, X. bemarivensis extends further into the dry forests of southwestern Madagascar than any other Xylopia .Specimens with flowers have been collected from October to February and in April, and with fruits from October to June. Seeds were reported to be eaten by the lemur Propithecus tattersalli ( Meyers 92 ), but no additional details were given. Xylopia bemarivensis has the second largest EOO and AOO of any Madagascar Xylopia species, 171 046 km 2 and 172 km 2 , respectively. One of the least likely Xylopia species in Madagascar to be of conservation concern, X. bemarivensis is nevertheless given a preliminary conservation assessment of Data Deficient because we have no information about habitat alteration or population structure or size. LOCAL NAMES. — Azoambo ( Bardot-Vaucoulon 915 ), fandriambarika ( Ramananjanahary 121 ), fandribarika ( Réserves Naturelles 3338 ), fotsiavadika ( Andrianantoanina et al. 848 ), fotsimavo ( Bardot-Vaucoulon et al. 1263 ), hazoambo ( Rakotondrafara et al. 430, Service Forestier (Comtet) SF 34395 ), hazoambolahy ( Hong-Wa 303 ), hazomavo ( Randrianarivelo et al. 159 ), marohefaka ( Manjakahery & Sola 72 ), moranga ( Ratovoson et al. 1045 ). DESCRIPTION Tree or shrub up to 20 m tall; d.b.h. up to 22 cm . Twigs short erect-pubescent, the hairs 0.2-0.3 mm long, eventually glabrate; nodes with one or occasionally two axillary branches. Leaves with larger blades 4.0- 9.8 cm long, 1.9-3.4 cm wide, chartaceous, concolorous or slightly paler abaxially, narrowly to broadly lanceolate, oblong, or elliptic, apex obtuse to rounded or occasionally acute, base broadly cuneate to obliquely rounded or subcordate, short-decurrent on petiole, margin flat, not revolute, pubescent, sometimes with hairs only along the midrib or glabrate adaxially, pubescent abaxially; midrib concolorous adaxially, secondary veins weakly brochidodromous, 9-16 per side, diverging at 45-80° from midrib, plane adaxially, slightly raised abaxially, higher-order veins indistinct to slightly raised on both surfaces; petiole 4.5-7 mm long, flat, smooth, pubescent. FIG. 16. — A , Fruit of X. australis , sp. nov. ( Rakotovao 4575 ); B , flower of X. bemarivensis Diels ( Nusbaumer 8139 ); C , flower bud of X. galokothamna , sp. nov. ( Ratovoson 2065 ); D , fruit of X. galokothamna , sp. nov. ( Ratovoson 2065 ); E , fruit of plant resembling X. galokothamna , sp. nov. , but with different pericarp pattern (see text) ( Buerki 20 ); F , monocarps and leafy shoots of X. lastelliana Baill. ( Rakotovao 4603 ); G , flowers of X. australis , sp. nov. ( Schatz 2641 ); H , monocarps of X. kalabenonensis D.M.Johnson, Deroin, & Callmander ( Razafitsalama et al. 1041 ) . Photos: A , F , C. Rakotovao; B , L. Nusbaumer; C , D , F. Ratovoson; E , M. W. Callmander; G , G. E. Schatz; H , J. Razafitsalama. A , C -H , used under the terms of the Creative Commons license for Tropicos, the botanical information system at the Missouri Botanical Garden; B , L. Nusbaumer (© Conservatoire et Jardin botaniques de la Ville de Genève). Inflorescences axillary or from the axils of fallen leaves on new flushes of growth, 1-flowered, not pedunculate, densely erect-pubescent; pedicels often obliquely bent near midpoint, 4.3-7.1 mm long, 0.5-1.0 mm thick; bracts 2, proximal bract attached near midpoint of pedicel and usually caducous, distal bract subtending sepals and persistent, 1.8-2.0 mm long, ovate to semicircular; buds narrowly oblong, apex obtuse. Sepals spreading at anthesis, 1/3-1/2-connate, 1.7-2.3 mm long, 2.4-2.8 mm wide, coriaceous, semicircular, apex broadly acute to obtuse, sericeous to tomentose abaxially. Petals white, cream-colored, or yellow, the inner petals also with a red blotch at the base in vivo ; outer petals spreading at anthesis, 9.7-12.2 mm long, 2.0- 3.3 mm wide at base, 1.2-1.8 mm wide at midpoint, coriaceous, narrowly lanceolate to lanceolate, keeled at apex adaxially, flat or faintly ridged abaxially, apex acute, densely puberulent except for glabrous base adaxially, densely sericeous except for glabrous base abaxially; inner petals more or less erect at anthesis, 7.3- 8.9 mm long, 1.6-2.4 mm wide at base, 0.7-1.0 mm wide at midpoint, coriaceous, linear-subulate, keeled on upper ¾ adaxially, keeled abaxially, apex acute, base concave with an undifferentiated margin, puberulent on both surfaces except for the sharply glabrous base. Stamens 100-120; fertile stamens 1.1-1.3 mm long, narrowly oblong, anther connective apex 0.2-0.3 mm long, shield-shaped to capitate, overhanging anther thecae, glabrous, anthers 9-11-locellate, filament 0.2-0.6 mm long; outer staminodes c. 1.1 mm long, oblong or broadly clavate, apex truncate; inner staminodes 0.9-1.1 mm long, oblong or clavate, apex truncate; staminal cone 1.3-2.0 mm in diameter, c. 0.8 mm high, completely concealing the ovaries, rim even or slightly laciniate. Carpels 3-5; ovaries c. 0.6 mm long, oblong, pubescent, stigmas loosely connivent, 1.4-1.8 mm long, filiform, with a few hairs at the apices. Fruit of up to 9 monocarps borne on a pedicel 6-9 mm long, 4.2-5 mm thick, glabrate; torus 7-11 mm in diameter, 6-7 mm high, globose to depressed-globose; monocarps with reddish green or purplish green to red exterior and red endocarp in vivo , 2.8-4.6 cm long, 1.3-2.0 cm wide, 1.3-1.6 cm thick, ellipsoid to oblong, weakly torulose, apex obtuse to rounded, base usually contracted into a stipe 3-5 mm long, 3-6 mm thick, verrucose, longitudinally wrinkled, sometimes also with a ridge along the abaxial surface, glabrate; pericarp 0.6- 1.2 mm thick. Seeds up to 7 per monocarp, in two rows, perpendicular to long axis, 9.7-14 mm long, 7.6-10 mm wide, 6.0- 6.6 mm thick, obovoid, oblong, or ellipsoid, irregularly elliptic in cross-section, brown to dark brown, smooth, dull, perichalazal ring not elevated; sarcotesta orange to red in vivo , sometimes visible as a white crust on dried seeds; aril absent. NOTES Xylopia bemarivensis is distinctive among Madagascar species of the genus for its pubescent twigs, thin-textured hairy leaves blunt at the apex and rounded at the base, one-flowered inflorescences with relatively long pedicels, and orange seeds. The only sympatric species of Xylopia in these dry forests is X. sericolampra , which has subcoriaceous leaves that are truncate at the base, silvery appressed hairs on the buds and perianth, flowers that emerge from leafless branches, and single pear-shaped monocarps. There is variation in flower color, with mature flowers varying from yellow-green on some specimens to white on others. Petal length is also variable, but in this case petal length can be variable within a single plant: the outer petals on the specimen Service Forestier 8066 SF range in length from 9.9 to 12.1 mm , or nearly the full range of variation seen for the species. The flowers are reported by many collectors to be fragrant, on the label of Gautier et al. LG 5964 as having an “odeur citronnée.” On the basis of its orange to red sarcotesta and thin pubescent leaves, Xylopia bemarivensis seems allied to the Xylopia odoratissima subgroup of section Stenoxylopia distributed across eastern and southern Africa ( Johnson & Murray 2018 ).