Xylopia anomala D.M.Johnson & N.A.Murray, 2020

Johnson, David M. & Murray, Nancy A., 2020, A revision of Xylopia L. (Annonaceae): the species of Madagascar and the Mascarene islands, Adansonia 42 (1), pp. 1-88 : 42-44

12. Xylopia anomala D.M.Johnson & N.A.Murray , sp. nov. ( Figs 6G ; 15A-K )

Belonging the Madagascar group of Xylopia species 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 cm long and 3.2 cm thick.

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.), Martial et 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.), Rakotonirina et 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.), Rakotonirina et al. 105 ( OWU, P[P01047882]) ; Sava Region, Sava, Andapa , Ambodivohitra , forêt Gelose à 1.5 km E du village de Marovato, 710 m, 14°34’40”S, 49°38’32”E, 16.II.2006 (buds), Rakotovao et al. 2680 ( MO, OWU, P[P01954121]) .

Prov. Toamasina, Masoala Peninsula, coastal trail leading S from Ambanizana , 0-10 m, 10.XII.1990 (fr.), Schatz & Modeste 3042 ( 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. 306 reports the fruit to be consumed by the Madagascar blue pigeon, Alectroenas madagascariensis. With an EOO and AOO estimated as 3468 km 2 and 24 km 2, 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 Tree up to 16 m tall; d.b.h. up to 12 cm.

Twigs finely appressed-pubescent, the hairs 0.1-0.3 mm long, at length glabrate; nodes with one or sometimes two axillary branches.

Leaves with larger blades 14.0-19.0 cm long, 5.3-6.5 cm wide, chartaceous, slightly discolorous, dull green adaxially, brown to orange-brown abaxially, elliptic, oblong, or oblanceolate, apex obtuse, acute, or acuminate, the acumen 18-22 mm long, 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 mm long, semi-terete, smooth, sparsely pubescent.

Inflorescences axillary, 2-5-flowered, corymb-like, pedunculate, pubescent; peduncles 2-3 mm long; pedicels 2-5 per peduncle, 5.4-6.2 mm long, 1.1-1.4 mm thick; bracts 3, evenly spaced along pedicel, caducous or distal bract persistent, 1.7-3.3 mm long, ovate to broadly ovate; buds oblong, apex rounded.

Sepals slightly spreading at anthesis, ¼-1/3-connate, 3.6-5 mm long, 3.7-4.8 mm wide, subcoriaceous, broadly ovate to triangular, apex obtuse, pubescent abaxially.

Petals light yellow in vivo; outer petals erect (ṙ) at anthesis, 8.6-9.0 mm long, 2.5-4.1 mm wide at base, 4.9-6.8 mm wide 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 mm wide at base, 2.5- 2.7 mm wide 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 mm long, narrowly oblong or slightly clavate, anther connective apex 0.1-0.3 mm long, shield-shaped, overhanging anther thecae, papillate, anthers 4-7-locellate, filament 0.4-0.6 mm long; outer staminodes 1.8-2.0 mm long, oblong to clavate, apex rounded to truncate; inner staminodes 1.1-1.4 mm long, oblong to clavate, apex truncate; staminal cone 1.6-2.4 mm in diameter, 0.7-1.0 mm high, concealing only the bases of the ovaries, rim irregularly laciniate.

Carpels c. 5; ovaries 1-1.4 mm long, narrowly oblong, densely pubescent, stigmas loosely connivent, 1.5-1.9 mm long, clavate and somewhat falciform, glabrous or with a few hairs at the apex.

Fruit of up to 4 monocarps borne on a pedicel 10-12 mm long, 5-6 mm thick, sparsely pubescent to glabrate; torus 11-15 mm in diameter, c. 7 mm high, depressed-globose; monocarps with light brown exterior in vivo, 5.8-7.3 cm long, 1.9-2.6 cm wide, 1.3-3.2 cm thick, oblong to oblanceolate, not torulose, apex obtuse, sometimes contracted into a short beak c. 3.5 mm long, base narrowed, sessile or contracted into a stipe c. 2 mm long, 8 mm thick, smooth, densely lenticellate, dull, glabrate or with scattered patches of pubescence; pericarp 1.5-2 mm thick.

Seeds c. 9 per monocarp, in two rows, perpendicular to long axis, 13.3-17.2 mm long, 9.0- 15.6 mm wide, 6.7-9.1 mm thick, ellipsoid to nearly globose, wedge-shaped to elliptic in cross-section, brown, smooth, dull, perichalazal ring visible but not elevated; sarcotesta color in vivo unknown, 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 Xylopia at 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 Xylopia species, and rare in the genus, although this fruit morphology is known in species such as X. hypolampra in Africa ( Johnson & Murray 2018) and X. ngii in 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 Stenoxylopia clade with species such as X. danguyella and X. fananehanensis , but without any close resemblance to other species within that group.

Figures

FIG. 6. — A, Shoot of X. amplexicaulis (Lam.) Baill., showing flower bud at left and clasping leaf bases visible at several nodes; B, branch of X. ravelonarivoi, sp. nov., with fruit (Ravelonarivo 4171); C, flower of X. humblotiana Baill., side view (Schatz 4475); D, buds of X. richardii Boivin ex Baill.; E, close-up of flower interior of X. lamarckii Baill., showing red coloration at bases of petals, and the stamens and stigmas; outer staminodes are visible along the edge of the androecium; F, flower of X. richardii at anthesis, one outer petal and one inner petal removed, so that androecium and stigmas are visible; G, leaves and fruit of X. anomala, sp. nov., (Rakotonirina 48); H, leaves and fruit of X. sericolampra Diels. Photos: A, Carlos Magdalena; B, D. Ravelonarivo; C, G. E. Schatz; D-F, C. Kaiser-Bunbury; G, C. Birkinshaw; H, R. N. Rabarijaona. B, C, G, used under the terms of the Creative Commons license for Tropicos, the botanical information system at the Missouri Botanical Garden; H, used under the terms of the the Creative Commons license for iNaturalist.

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.

FIG. 28. — Distributions of Xylopia ambanjensis Cavaco & Keraudren, X. anomala, sp. nov., X. australis, sp.nov., X.beananensis Cavaco & Keraudren, X. bemarivensis Diels,X.buxifolia Baill., X.capuronii Cavaco & Keraudren,X. carinata, sp. nov., X. danguyella Ghesq. ex Cavaco & Keraudren, X. dielsii Cavaco & Keraudren, X. fananehanensis Cavaco & Keraudren, and X. flexuosa Diels on Madagascar.