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