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
43
42
Tracheophyta
species
anomala
sp. nov.
( Figs 6G; 15A-K)
Belonging the Madagascargroup of Xylopiaspecies having short broad petals and clavate stigmas, but distinguished from the others by the chartaceous leaves lacking a pronounced vein reticulum, the flowers up to 5 per inflorescence and forming a short corymb-like inflorescence, the obtuse outer petals 8.6-9.0 mm long and 2.5-4.1 wide at base, the stamens only 4-7-locellate, the staminal cone with filament bases only loosely connate, and the densely lenticellate monocarps up to 7.3 cmlong and 3.2 cmthick.
TYPE. — Madagascar. Prov. Antsiranana, Sava Region, Anjangoveratra, Antanandava, Makirovana, 14°10’01”S, 49°57’12”E, 685 m, 4.V.2010(fl.), Razakamalala et al. 5422(holo-, MO!). PARATYPES. — Madagascar. Prov. Antsiranana, Sava Region, Vohemar, Antsirabe-Nord, Antanambaon’Isokitra, forêt de Tsihomanaomby, au sud-est du village Antanambaon’Isokitra, 14°06’S, 50°02’E, 30.XI.2013(fr.), Martialet al. 306( OWU, P[P00982778]); Sava Region, Vohemar, Antsirabe Nord, Andravinambo, forêt d’Antsolatra, 14°06’45”S, 49°57’44”E, 387 m, 27.II.2013(fr.), Rakotonirinaet al. 48( OWU, P[P01030032]); Sava Region, Sambava, Marogaona, Ambodivapaza, forêt de Farahangitra, 14°08’09”S, 49°54’43”E, 322 m, 11.IV.2013(fl.), Rakotonirinaet al. 105( OWU, P[P01047882]); Sava Region, Sava, Andapa, Ambodivohitra, forêt Geloseà 1.5 kmE du village de Marovato, 710 m, 14°34’40”S, 49°38’32”E, 16.II.2006(buds), Rakotovaoet al. 2680( MO, OWU, P[P01954121]). Prov. Toamasina, Masoala Peninsula, coastal trail leading S from Ambanizana, 0-10 m, 10.XII.1990(fr.), Schatz & Modeste3042( K, MO, P[ P01966122]).
DISTRIBUTION, ECOLOGY AND CONSERVATION STATUS. — Xylopia anomala, sp. nov., is restricted to northeastern Madagascar, occurring in humid forest at elevations from near sea level up to nearly 700 meters ( Fig. 28); one collection label mentions a blackish soil substrate and another sand. Specimens with large buds or flowers have been collected in February, April, and May, and with fruits in February, November, and December. The label of Martial et al. 306reports the fruit to be consumed by the Madagascarblue pigeon, Alectroenas madagascariensis. With an EOO and AOO estimated as 3468 km2 and 24 km2, respectively, Xylopia anomala, sp. nov., is given a preliminary conservation assessment of Endangered ( Table 2).
LOCAL NAME. — Moranga ( Rakotonirina et al. 48, Martial et al. 306).
DESCRIPTION Treeup to 16 mtall; d.b.h. up to 12 cm. Twigsfinely appressed-pubescent, the hairs 0.1-0.3 mmlong, at length glabrate; nodes with one or sometimes two axillary branches. Leaveswith larger blades 14.0-19.0 cm long, 5.3-6.5 cmwide, chartaceous, slightly discolorous, dull green adaxially, brown to orange-brown abaxially, elliptic, oblong, or oblanceolate, apex obtuse, acute, or acuminate, the acumen 18-22 mmlong, base cuneate, decurrent on petiole, margin flat, not revolute, glabrous adaxially, glabrous or with a few scattered hairs along the midrib abaxially; midrib dull yellow or blackish brown toward base adaxially, secondary veins weakly brochidodromous, 8-15 per side, diverging at 40-70° from midrib, these and higher-order veins plane to raised adaxially, raised abaxially; petiole 4.5-13 mmlong, semi-terete, smooth, sparsely pubescent. Inflorescencesaxillary, 2-5-flowered, corymb-like, pedunculate, pubescent; peduncles 2-3 mmlong; pedicels 2-5 per peduncle, 5.4-6.2 mmlong, 1.1-1.4 mmthick; bracts 3, evenly spaced along pedicel, caducous or distal bract persistent, 1.7-3.3 mmlong, ovate to broadly ovate; buds oblong, apex rounded. Sepalsslightly spreading at anthesis, ¼-1/3-connate, 3.6-5 mmlong, 3.7-4.8 mmwide, subcoriaceous, broadly ovate to triangular, apex obtuse, pubescent abaxially. Petalslight yellow in vivo; outer petals erect (ṙ) at anthesis, 8.6-9.0 mm long, 2.5-4.1 mmwide at base, 4.9-6.8 mmwide at midpoint, fleshy, ovate, concave, keeled at the apex adaxially, flat or slightly ridged abaxially, apex obtuse, densely pubescent except for the glabrous base adaxially, pubescent abaxially; inner petals erect and possibly connivent at base at anthesis, 8.2-9.0 mm long, 2.5-3.3 mmwide at base, 2.5- 2.7 mmwide at midpoint, fleshy, lanceolate-oblong, keeled at the apex adaxially, keeled abaxially, apex broadly acute to obtuse, base shallowly concave into a short claw, with undifferentiated margin but with a transverse thickening above the concavity, densely hairy except for the glabrous base on both surfaces. Stamens c.100; fertile stamens 1.4-1.7 mmlong, narrowly oblong or slightly clavate, anther connective apex 0.1-0.3 mmlong, shield-shaped, overhanging anther thecae, papillate, anthers 4-7-locellate, filament 0.4-0.6 mmlong; outer staminodes 1.8-2.0 mm long, oblong to clavate, apex rounded to truncate; inner staminodes 1.1-1.4 mmlong, oblong to clavate, apex truncate; staminal cone 1.6-2.4 mmin diameter, 0.7-1.0 mm high, concealing only the bases of the ovaries, rim irregularly laciniate. Carpels c.5; ovaries 1-1.4 mmlong, narrowly oblong, densely pubescent, stigmas loosely connivent, 1.5-1.9 mmlong, clavate and somewhat falciform, glabrous or with a few hairs at the apex. Fruitof up to 4 monocarps borne on a pedicel 10-12 mmlong, 5-6 mmthick, sparsely pubescent to glabrate; torus 11-15 mmin diameter, c. 7 mmhigh, depressed-globose; monocarps with light brown exterior in vivo, 5.8-7.3 cmlong, 1.9-2.6 cmwide, 1.3-3.2 cmthick, oblong to oblanceolate, not torulose, apex obtuse, sometimes contracted into a short beak c. 3.5 mmlong, base narrowed, sessile or contracted into a stipe c. 2 mmlong, 8 mmthick, smooth, densely lenticellate, dull, glabrate or with scattered patches of pubescence; pericarp 1.5-2 mmthick. FIG. 15. — Xylopia anomala, sp. nov.and X. australis, sp. nov.: A -K, X. anomala, sp. nov.; A, habit; B, seed, view from micropylar end; C, seed, side view; D, fruit; E, inflorescence; F, inflorescence with mature flower; G, inner petal, adaxial view; H, outer petal, adaxial view; I, staminal cone, ovaries, and outer staminodes; J, carpel; K, fertile stamen, abaxial view. L -U, X. australis, sp. nov.: L, outer staminode, abaxial view; M, fertile stamen, abaxial view; N, inflorescence; O, stigma; P, inner petal, adaxial view; Q, outer petal, adaxial view; R, habit; S, fruit; T, seed, view from micropylar end; U, seed, side view. A, from Rakotonirina 48(OWU); B -D, from Schatz & Modeste 3042(MO); E, from Rakotovao et al. 2680(MO); F -I, K, from Razakamalala et al. 5422(MO); J, from Rakotovao et al. 2680(OWU); L, M, O -Q, from Schatz 2641(MO); N, from Rakotovao et al. 4575(MO); R -U, from Razakamalala et al. 2414(MO). Scale bars: A-D, R-U, 1 cm; E, F, G, H, N, P, Q, 5 mm; I-M, O, 1 mm. Seeds c.9 per monocarp, in two rows, perpendicular to long axis, 13.3-17.2 mmlong, 9.0- 15.6 mmwide, 6.7-9.1 mmthick, ellipsoid to nearly globose, wedge-shaped to elliptic in cross-section, brown, smooth, dull, perichalazal ring visible but not elevated; sarcotesta color in vivounknown, visible as white crusty patches on dried seeds; aril absent.
NOTES The first impression of Xylopia anomala, sp. nov., both in flower and in fruit, is that it is not a Xylopiaat all. The leaves are large, papery, and broad, the flower buds are rounded rather than pointed, and the inflorescences are corymb-like. Densely lenticellate oblong woody monocarps are unique among Madagascar Xylopiaspecies, and rare in the genus, although this fruit morphology is known in species such as X. hypolamprain Africa ( Johnson & Murray 2018) and X. ngiiin Asia ( Johnson & Murray 2015). Unusual features of the plant extend to the androecium: the stamens have a low number of anther locelli, 4-7, and are borne on a staminal cone in which the stamen filaments are only loosely connate ( Fig. 15I). The number of characters unusual for the genus suggested the specific epithet. The short, broad, and thick petals, together with the clavate stigmas, place this species in the subclade of the Stenoxylopiaclade with species such as X. danguyellaand X. fananehanensis, but without any close resemblance to other species within that group.
2632071769
2010-05-04
MO
Madagascar
685
-14.1669445
Anjangoveratra
21
49.953335
Sava Region
43
42
1
Prov. Antsiranana
holotype
2632071366
2013-11-30
OWU
Antanambaon'Isokitra & de Tsihomanaomby & Martial
Madagascar
Sava Region
-14.1
Antsirabe-Nord
1289
50.033333
Vohemar
43
42
1
Antsiranana
paratype
2632071690
2013-02-27
OWU
Rakotonirina
Madagascar
Sava Region
387
-14.1125
Antsirabe Nord
21
49.962223
Vohemar
43
42
1
Prov.
paratype
2632071414
2013-04-11
OWU
de Farahangitra & Rakotonirina
Madagascar
Sava Region
322
-14.135833
Marogaona
21
49.911945
Sambava
43
42
1
Prov.
paratype
2632071560
2006-02-16
MO, OWU
de Marovato & Rakotovao
Madagascar
Sava Region
710
-14.577778
Andapa
21
49.642223
Sava
43
42
1
Prov.
paratype
2632071372
1990-12-10
K, MO, P
Schatz & Modeste
5
coastal trail leading S from Ambanizana
Masoala Peninsula
43
42
P01966122
1
Prov. Toamasina
paratype