A revision of Xylopia L. (Annonaceae): the species of Madagascar and the Mascarene islands Johnson, David M. Murray, Nancy A. Adansonia 2020 2020-02-05 42 1 1 88 D.M.Johnson & N.A.Murray D. M. Johnson & N. A. Murray 2020 Magnoliopsida Annonaceae Xylopia Plantae Magnoliales 33 32 Tracheophyta species ravelonarivoi sp. nov. ( Figs 6B; 11)  Species resembling  Xylopia humblotianaBaill.in its small blunt coriaceous leaves, obtuse petals, and narrowly oblong monocarps, but differing in the tree habit, more strongly decurrent leaf bases, densely pubescent pedicels and sepals, lanceolate outer petals, and more strongly wrinkled monocarps borne on pedicels 13-19 mmlong and 1.6-3.7 mmthick.  TYPE. —   Madagascar. Prov. Toamasina, Antsimanana Region: Brickaville, Maroseranana, Ambodilendemy, suivant la rivière d’Andrangato, 18°26’05”S, 48°46’51”E,  658 m,  14.III.2011(fl.), Ravelonarivo et al. 3641(holo-, MO!).  PARATYPES. —   Madagascar. Prov. Antsiranana, Sava Region, Diego, Sambava, Anjangoveratra, Anamboafo, forêt de Makirovana-Beraharaha, 14°09’07”S, 49°57’13”E,  549 m,  6.III.2014(fl.),  Martialet al. 548( MO);  Sava Region, Sambava, Anjangoveratra, Anamboafo, forêt de Belalitra, Makirovana, 14°09’14”S, 49°58’09”E,  585 m,  8.III.2014(fl.),  Rakotonirinaet al. 537( MO);  MarojejyRNI, Analamboahangy, Andrakata, Andapa, aux environs de Manenobasy, 14°35’16”S, 49°41’20”E,  1171 m,  18-24.I.1995(fr.),  Rasoavimbahoaka509( MO, P[P01954048]);  Andrakata Canton, Andapa District[ 14°37’S, 49°43’E],  13.VIII.1957(fr.),  Safy9089 RN(P[P01954046]).  Prov. Mahajanga, Ruisseau Ambatoharanana Andranomena commune rurale de Matsoandakana, district de Befandrina Nord, 15°08’16”S, 49°21’02”E,  1045 m,  12.II.2008(fr.), Bernard et al. 778( K, MO).  Prov. Toamasina, Fivondronana Maroantsetra, Commune Ambinanitelo, Fokontany Marovovonana, Ankiriandro Mountain, 15°17’37”S, 49°33’10”E,  640 m,  10.I.2003(fr.), Antilahimena et al. 1683( K, P[ P01954047]);  Analanjirofo Region, Fivondronana Vavatenina, Commune Ambodimangavalo, à 3 kmà l’Ouest du village Ambatoharanana, dans le Parcnational de Zahamena, le long de la piste vers Antevibe, 17°33’54”S, 48°53’18”E,  875 m,  29.IV.2003(fr.), Rakotonandrasana 676( OWU);  Toamasinasous-préfecture, Vavatenina, commune: Ambodimangavelo, Moango, dans l’Aire protégée de Zahamena, 17°33’32”S, 48°53’55”E,  800 m,  4.V.2003(immature fr.), Rakotondrajaona et al. 281(K, MO);  Antsinanana Region, Vohimanana, Dist. Brickaville, Com. Maroseranana, Fkt. Ambatolampy, 18°23’27”S, 48°48’11”E,  846 m,  23.I.2012(fr.), Ravelonarivo et al. 4171( MO).  DISTRIBUTION, ECOLOGY AND CONSERVATION STATUS. —  Xylopia ravelonarivoi, sp. nov., occurs in humid lowland and lower montane forest on ridges and slopes in northeastern Madagascarat elevations from 549 to 1171 meters.Specimens have been collected with flowers in March and with fruits in January, February, April, and May. With an EOO and AOO estimated as 7118 km 2and 40 km 2, respectively, the species is given a preliminary conservation assessment of Endangered. It is documented by two collections from within the protected area of Zahamena National Park and one collection from the Marojejy Reserve.  LOCAL NAMES. — Hazoambo ( Rakotondrajaona et al. 281), hazoambo boriboriravina ( Rakotonandrasana 676), moranga ( Martial et al. 548), morango ( Rakotonirina et al. 537).  DESCRIPTION  Treeup to 15 mtall, or rarely a shrub up to 8 m; d.b.h. up to 30 cm.  Twigsdensely appressed-pubescent, the hairs 0.1-0.3 mmlong, eventually glabrate; nodes with one axillary branch.  Leaveswith larger blades 5.0- 7.2 cmlong, 2.0- 3.4 cmwide, subcoriaceous to coriaceous, discolorous, dark brown to gray and often shiny adaxially, light brown and dull abaxially, obovate, oblanceolate, elliptic, or elliptic-oblong, apex obtuse, rounded, retuse, or emarginate, base cuneate, decurrent on petiole, margin slightly recurved or revolute, glabrous adaxially, sparsely appressed-pubescent, especially along midrib, abaxially; midrib dull red to brown, darkening toward base adaxially, secondary veins weakly brochidodromous, 8-11 per side, diverging at 50-70° from midrib, these and higherorder veins raised and forming a conspicuous reticulum on both surfaces; petiole 1.5-7 mmlong, shallowly canaliculate or a little flattened, smooth or faintly wrinkled, glabrous or with a few hairs.  Inflorescencesaxillary from the axils of fallen leaves, 1-3-flowered, pedunculate, densely appressed-pubescent; peduncle 1.7-1.8 mmlong; pedicels up to 3 per peduncle, 4.8-9.7 mmlong, 1.0- 1.2 mmthick; bracts 2, one near base of the pedicel and the other near midpoint, caducous, only scars seen; buds lanceolate, apex obtuse.  Sepalsspreading at anthesis, ¼-1/2-connate, 1.4-3.6 mmlong, 3.5-4 mmwide, coriaceous, broadly triangular, apex acute or rarely obtuse, appressed-pubescent abaxially.  Petalsgreenish white in vivo; outer petals slightly spreading at anthesis, 10.0- 12.2 mmlong, 2.7-4.5 mmwide at base, 2.4-3.0 mm wide at midpoint, fleshy, lanceolate, keeled at the apex but otherwise flat adaxially, flat abaxially, apex obtuse, densely pubescent except for glabrous base adaxially, densely appressed-pubescent abaxially; inner petals slightly spreading at anthesis, 10.5-10.9 mmlong, 2.2-3.0 mm wide at base, 0.9-1.2 mmwide at midpoint, fleshy, linear-subulate, keeled on both surfaces, apex acute, concave with undifferentiated margin, densely pubescent on both surfaces except for glabrous base, with a band of papillae across widest point adaxially.  FIG. 11. —  Xylopia ravelonarivoi, sp. nov.: A, habit; B, leaf base,oblique adaxial view to show decurrence of leaf blade on petiole; C, flower bud, side view; D, leaf; E, fruit; F, fertile stamen, abaxial view; G, outer staminode, abaxial view; H, adaxial staminode, abaxial view; I, outer petal, adaxial view; J, inner petal, adaxial view; K, seed, side view; L, seed, view of micropylar end. A, from Rakotondrajaona et al. 281(MO); B, D, E, K, L, from Ravelonarivo et al. 4171(MO); C, F -J, from Ravelonarivo et al. 3641(MO). Scale bars: A, D, E, 1 cm; B, C, I-L, 5 mm; F-H, 1 mm.  Stamens c.250; fertile stamens 1.1-1.4 mmlong, narrowly oblong, anther connective apex 0.2-0.3 mmlong, shield-shaped to capitate, overhanging anther thecae, minutely papillate, anthers 7-9-locellate, filament 0.4-0.6 mmlong; outer staminodes c. 1.3 mmlong, oblong to clavate, apex truncate or rounded; inner staminodes 1.0- 1.1 mmlong, clavate, apex truncate; staminal cone 2.2-2.6 mmin diameter, 1.0- 1.5 mmhigh, completely concealing the ovaries, rim even.  Carpels10-13; ovaries 1.0- 1.1 mmlong, narrowly oblong, pubescent, stigmas connivent, 1.8-2.2 mmlong, filiform, glabrous or with a few apical hairs.  Fruitof up to 13 monocarps borne on a pedicel 13-19 mmlong, 1.6-3.7 mmthick, glabrate or with a few scattered hairs; torus 5-12 mmin diameter, 3.5-9 mmhigh, depressed-globose; monocarps with red exterior in vivo, 2.2-4.5 cmlong, 0.5-0.9 cmwide, 0.5-0.8 cmthick, narrowly oblong, often slightly torulose, apex obtuse to rounded, sometimes slightly beaked, base tapered gradually, sometimes forming a stipe c. 4 mmlong, 3 mmthick, obliquely wrinkled, shiny, glabrate; pericarp 0.3-0.8 mmthick.  Seedsup to 5 per monocarp, in a single row, nearly parallel to long axis, 5.6-7.8 mmlong, 2.9-4.6 mmwide, 3.3-4.0 mm thick, broadly ellipsoid to oblong, elliptic in cross-section, dark brown to black, smooth or slightly wrinkled and pitted, shiny, perichalazal ring slightly raised; sarcotesta absent; aril white in vivo, white to amber-colored when dried, bilobed, lobes 2.0- 2.5 mmlong, 3.0- 3.6 mmwide, fleshy, surface wrinkled.  NOTES   Xylopia ravelonarivoi, sp. nov., is similar to  X. humblotianabut differs in a number of characters that, together with the ecological differences, distinguish the two species. It is usually a tree up to 15 mtall, rather than a shrub or treelet. The leaves have a more decurrent base, such that the demarcation between blade and petiole is indistinct ( Fig. 11B). The pedicels and sepals are densely pubescent. The sepals are longer, and the outer petals are lanceolate rather than oblong; the latter character is also expressed in the buds that are lanceolate rather than oblong as in  X. humblotiana. The monocarps of this species are obliquely wrinkled, while those of  X. humblotianaare usually smooth. There is a sharp difference in elevation range between the two species:  X. ravelonarivoi, sp. nov., grows at elevations from 550 to 1170 mand  X. humblotianafrom sea level to  50 m.  Xylopia madagascariensisalso has monocarp and leaf sizes similar to those of  X. ravelonarivoi, sp. nov., but its leaves are blunt- acuminate rather than obtuse to rounded and its monocarps are smooth and finely verrucose rather than strongly wrinkled. In addition,  X. madagascariensisreaches its upper elevational limit at just over 500 meters, while the lower elevational limit for  X. ravelonarivoi, sp. nov., is c.550 meters. The specimen Rakotondrajaona et al. 281was the source of molecular data used in the phylogenetic analysis of  Xylopiaby Stull et al.(2017), where it was identified as  X. humblotiana; the seed illustrated in Figure 4Hof that work is, however, correctly identified as  X. humblotiana. The species is named for Désiré Ravelonarivo, the collector of the typespecimen, who has contributed to a clearer understanding of  Xylopiadiversity on Madagascarby making 16 critical  Xylopiacollections representing six species. 2632071780 2011-03-14 MO Madagascar 658 -18.434721 Brickaville 21 48.780834 Antsimanana Region 33 32 1 Prov. Toamasina holotype 2632071413 2014-03-06 MO Anamboafo & de Makirovana-Beraharaha & Martial Madagascar Prov. 549 -14.151944 Diego 21 49.953613 Sava Region 33 32 1 Antsiranana paratype 2632071463 2014-03-08 MO de Belalitra & Makirovana & Rakotonirina Madagascar Sava Region 585 -14.153889 Anjangoveratra 21 49.969166 Sambava 33 32 1 paratype 2632071789 1995-01-18 1995-01-24 1995-01-18 MO de Manenobasy & Rasoavimbahoaka Madagascar Marojejy 1171 -14.587777 Andrakata 21 49.68889 Analamboahangy 33 32 1 paratype 2632071418 1957-08-13 Safy Madagascar -14.616667 Andapa District 1287 49.716667 Andrakata Canton 33 32 1 paratype 2632071325 2008-02-12 K, MO 1045 -15.137777 commune rurale de Matsoandakana 21 49.350555 Ruisseau Ambatoharanana Andranomena 33 32 1 Mahajanga paratype 2632071509 2003-01-10 K, P 640 -15.293612 Commune Ambinanitelo 21 49.552776 Fivondronana Maroantsetra 33 32 P01954047 1 Prov. Toamasina paratype 2632071774 2003-04-29 OWU le Parc & de Zahamena 875 -17.564999 Analanjirofo Region 21 48.888336 33 32 1 paratype 2632071685 2003-05-04 MO de Zahamena 800 -17.558887 Toamasina 21 48.898613 33 32 1 Toamasina paratype 2632071482 2012-01-23 MO 846 -18.390833 Antsinanana Region 21 48.803055 33 32 1 Toamasina paratype