Xylopia lastelliana Baill.

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 : 67-68

26. Xylopia lastelliana Baill.

( Figs 1C ; 14Q-Y ; 16F )

Adansonia; Recueil d’Observations botaniques 4: 144 (1864). — Type: Madagascar. Without definite locality, 1841 (fl.), Lastelle s. n. (holo-, P[P030304]!) .

Xylopia lastelliana var. acuta Ghesq. ex Cavaco & Keraudren , Bulletin du Jardin botanique de l’État à Bruxelles 27: 88 (1957). — Type: Madagascar. Prov. Antsiranana [“ Est ”], Antalaha , IX.1912 (fl.), Perrier de la Bâthie 4940 (holo-, P [ P030405 ]!; iso-, B!) .

MATERIAL EXAMINED. — Madagascar. Prov. Antsiranana. Parc national de Masoala, c. 1.0- 1.5 km E of campsite at Ambatoledama,

720-792 m, 14.XII.1998 (fl., fr.), Flores & Andriantiana 72 (A, K n.v., MO); Marosena, Ampanavohana, 2.I.1955 (fl.), Réserves Naturelles de Madagascar 6953 ( OWU, P[P01987013, P01987014, P01987015, P02133044, P02133045, P02133046]) ; Sarahandrano- Antalaha, 8.IX.1958 (fr.), Service Forestier 19181 (K) .

Prov. Fianarantsoa. Bassin de Mananjary, entre Vatovavy et Mananjary, VIII.1911 (fl.), Perrier de la Bâthie 4944 (B, P[P01987033, P01987034]).

Prov. Toamasina. Environs de la Baie d’Antongil colline d’Ambodiatafana, au NW de l’embouchure de la Rantabe, 12.IX.1957 (fr.), Capuron 18258 SF (BR, K n.v., P[P01986998, P01986999]); Fivondronana : Ambodiriana , Firaisina : Fontsimavo , Réserve naturelle intégrale de Betampona, 17°54’46”S, 49°12’20”E, 427 m, 20.XI.2001 (bud, fr.), Rabehevitra et al. 24 ( MO, P[P06839267]), 20.XI.2001 (fl.), Rabehevitra et al. 25 ( MO, P[P06839285]) ; Atsinanana Region , Atsinanana , Vatomandry , Ambalabe , Sahanionaka, route Vodiriana–Ambalabe, 19°11’31”S, 48°34’40”E, 305 m, 19.IX.2011 (fr.), Rakotoarivelo et al. 558 ( MO, P[P00903563]) ; Analanjirofo Region , Masoala Parc national, Ambatolaidama , Mahalevona, Maroantsetra, 15°17’21”S, 50°00’08”E, 0-1000 m, 1.X.1997 (fr.), Ranaivojaona et al. 173 ( MO, OWU, P[P01986997]) ; Atsinanana Region , Commune : Ambodiriana , Fokontany : Fontsimavo ( Fotsimavo by map) ( Andrantabe ), Andrendrirendry, Réserve naturelle intégrale de Betampona, 17°55’46”S, 49°12’14”E, 341 m, 20.XI.2001 (fr.), Razafitsalama 119 ( MO, P[P00903551]) ; Atsinanana Region , Atsinanana , Vatomandry, Ambalabe, Ambinanindrano II, forêt de Vohibe, 19°10’17”S, 48°32’05”E, 548 m, 16.VIII.2014 (fr.), Razanatsima et al. 1320 ( MO, P) ; Masoala Peninsula just S of Ambanizana along coastal trail and just E of Ambanizana, 15°38’S, 49°58’E, 70 m, 21.XII.1989 (fl.), Schatz & Modeste 2877 ( BR, K n.v., MO, P[P01987012], WAG) ; Masoala Peninsula , “South Trail” (S of Androka River), climbing hills SE of Ambanizana, 15°38’S, 49°59’E, 250 m, 24.XII.1990 (fl.), Schatz & Modeste 3086 (G, K n.v., MO, PRE, WAG) ; F. Amelamena, Vohilengo , Soanierana-Ivongo, 23.VII.1954 (fr.), Service Forestier 11062 ( OWU, P[P01987000, P01987001, P01987002]) ; Ampasina , District Fénérive-Est, s. d. (fl.), Service Forestier 19155 (K) .

Prov. Toliara. Commune rurale Mahatalaky, Fokontany Tsialagna, village proche Tsialagna, 24°43’01”S, 47°00’39”E, 190 m, 19.XI.2009 (fl. buds, fr.), Rakotovao et al. 4603 (MO); Anosy Region , District Fort-Dauphin , Commune Ranomafana , Fokontany Magatsika, forêt Tsirandrany, 24°35’06”S, 47°02’14”E, 669 m, 21.XI.2009 (fr.), Razakamalala 4755 ( MO) .

DISTRIBUTION, ECOLOGY AND CONSERVATION STATUS. — Restricted to humid evergreen forests and occasionally littoral forests of northeastern Madagascar, Xylopia lastelliana usually occurs at elevations from 70 up to nearly 800 meters ( Fig. 29 ); two atypical specimens, discussed below, were collected at c. 1300 meters. Specimens with flowers have been collected in January, August, November, and December, and with fruits in February and from July to December. Xylopia lastelliana has the fourth largest EOO of any Madagascar species, 124 820 km 2, but a relatively small AOO of 72 km 2, and is given a preliminary conservation assessment of Data Deficient ( Table 2 ).

LOCAL NAMES. — Hazoambo (Rakotoarivelo et al. 558, Razafitsalama 119), moranga (Schatz & Modeste 2877, 3086).

DESCRIPTION Tree up to 25 m tall; d.b.h. up to 55 cm.

Twigs densely sericeous, the hairs 0.4-1.0 mm long, eventually glabrate; nodes with one or often two axillary branches.

Leaves with larger blades 3.5-6.7 cm long, 1.1-2.5 cm wide, subcoriaceous or chartaceous, concolorous, lanceolate, ovate, or elliptic, apex acuminate, the acumen 7-10 mm long, to acute, base cuneate to rounded, sometimes oblique, decurrent on petiole, margin flat, not revolute, glabrous or with a few hairs on the midrib adaxially, sericeous abaxially with silvery hairs; midrib slightly darkened toward base adaxially, secondary veins weakly brochidodromous, 8-13 per side, diverging at 50-60° from midrib, these and higher-order veins indistinct to slightly raised on both surfaces; petiole 1.5-3.5 mm long, shallowly canaliculate, smooth or slightly wrinkled, pubescent.

Inflorescences axillary, 1-4-flowered, pedunculate or not, densely rusty-pubescent; peduncle 0.6-5 mm long; pedicels 2 per peduncle or arising separately from leaf axil, 3.5-6.8 mm long, 0.6-1.0 mm thick; bracts 2, one attached near the pedicel midpoint and the other just proximal to the sepals, caducous or the distal bract sometimes persistent, 1.2-1.9 mm long, broadly ovate; buds lanceolate, apex acute, sometimes slightly falciform.

Sepals spreading at anthesis, ¹⁄5-1/2-connate, 1.8-2.5 mm long, 2.0- 2.5 mm wide, coriaceous, ovate to broadly triangular, apex acute, densely sericeous abaxially.

Petals yellow, cream-colored, or amber colored and the inner petals tinged with red at the base in vivo; outer petals erect at anthesis, 9.5-15.5 mm long, 2.0- 3.3 mm wide at base, 1.0- 1.7 mm wide at midpoint, coriaceous or slightly fleshy, linear to linear-lanceolate, flat adaxially, faintly keeled abaxially, apex obtuse, pubescent except for glabrous concave base adaxially, sericeous abaxially; inner petals bent or curved outward at anthesis, 8.5-11.2 mm long, 1.4-2.7 mm wide at base, 0.5-0.8 mm wide at midpoint, coriaceous or slightly fleshy, linear-subulate, keeled on both surfaces, apex obtuse, base concave with undifferentiated margin, densely puberulent except for the glabrous base on both surfaces.

Stamens c. 75; fertile stamens 0.8-1.2 mm long, narrowly oblong, anther connective apex 0.1-0.2 mm long, shield-shaped, overhanging anther thecae, papillate, anthers 7-10-locellate, filament 0.2-0.3 mm long; outer staminodes 1.0- 1.2 mm long, narrowly oblong to clavate, apex obtuse to rounded; inner staminodes 0.7-0.8 mm long, clavate, apex truncate or slightly emarginate; staminal cone 0.9-1.5 mm in diameter, 0.7-1.0 mm high, concealing the basal half of the ovaries, rim irregularly laciniate.

Carpels 3-5; ovaries c. 0.8 mm long, oblong, densely sericeous, stigmas connivent, 1.6-2.5 mm long, filiform, glabrous or occasionally pubescent.

Fruit of up to 6 monocarps borne on a pedicel 6-8 mm long, 4-5 mm thick, glabrate; torus 10-18 mm in diameter, 12- 14 mm high, depressed-globose to pyramidal; monocarps with greenish red, red, or brown exterior and pinkish red endocarp in vivo, 2.3-4.1 cm long, 1.8-2.5 cm wide, 1.8-2.7 cm thick, ellipsoid to ovoid, not torulose, apex rounded, base sessile or occasionally contracted into a stipe c. 4 mm long and 8 mm thick, sometimes faintly verrucose, slightly wrinkled, dull, lenticellate, glabrate; pericarp 1.4-1.8 mm thick.

Seeds up to 8 per monocarp, in two slightly overlapping rows, perpendicular to long axis, 10.8-15.2 mm long, 8.8-12.0 mm wide, 3.9-5.9 mm thick, oblong to nearly circular and narrowed into a short neck around the micropyle, narrowly oblong in cross-section, brown to reddish brown, smooth or slightly wrinkled, dull or slightly shiny, perichalazal ring sometimes slightly raised; sarcotesta color in vivo unknown, but sometimes visible as white crust on dried seeds; aril absent.

NOTES

Xylopia lastelliana is readily distinguished from most other Madagascar species of Xylopia by the densely and persistently sericeous abaxial leaf surfaces, the hairs most persistent along the midrib. Only X. sclerophylla , sp. nov., has similar leaf indument, and its leaves are smaller, acute to more often obtuse at the apex rather than acute to acuminate, and its monocarps are longitudinally ridged and only sparsely lenticellate. The monocarps of X. lastelliana are large and thick-walled, containing somewhat flattened seeds narrowed to a donutshaped ring around the micropyle similar to that seen in the seeds of X. buxifolia .

The variety distinguished by Cavaco and Keraudren, X. lastelliana var. acuta , merges imperceptibly into the nominate variety and is not recognized here; there is variation in shape of the leaf apex even on the same specimen in this species. Some specimens previously determined by Keraudren as X. lastelliana , however, are here segregated as Xylopia sclerophylla , sp. nov., which has leaves that are consistently less sericeous and more coriaceous, and monocarps with a thinner pericarp.

Two collections from the Maevarano River region of northwestern Madagascar, Capuron 3004 (OWU, P[P01989230]), Gorges de la Maevarano [c. 14°38’S, 48°17’E], 1350 m, 12.II.1950 (fr.), and Humbert & Capuron 25381 (OWU, P[P02034869]), Montagnes au Nord de Mangindrano (Haute Maevarano), jusqu’aux sommets d’Ambohimirahavavy (partage des eaux Mahavavy-Androranga: Centre Nord), vallon d’Ambatohafo [14°10’S, 48°58’E], 1350 m, 19.I.-12.II.1951 (fr.), while having leaves resembling those of X. lastelliana in shape and indument, are distant from other localities for the species and were collected from much higher elevations. Neither collection has flowers. The leaves are toward the small end of the range for X. lastelliana , 4.2-4.6 cm long and 1.5-1.6 cm wide. The monocarps are more oblong than ovoid or ellipsoid, c. 4.6 cm long, 2.7 cm wide, and 2.4 cm thick, wrinkled, and lacking lenticels, and have an even thicker pericarp (c. 2.6 mm), and the seeds do not have the donut-shaped ring around the micropyle seen in the seeds of X. lastelliana elsewhere. Similarly, the specimen Gautier et al. LG 3161 (K, P[P01954149]), from Réserve Spéciale de Manongarivo, Ambongomirahavavy (13°59’S, 48°17’E), confluence Manongarivo/Marofotrotro, 100 m, 29.IX.1996 (buds, immature fr.), has a silvery indument similar to that seen in X. lastelliana , and is from a tree 25 m tall with a d.b.h. of 40 cm, but its monocarps are smaller than those in both the Maevarano and eastern lowland material.

No locality is given in the protologue or on the label of the type specimen of X. lastelliana collected by de Lastelle, but given that de Lastelle made his home in Tamatave (now Toamasina; Ellis 1859) it is likely that the specimen came from the vicinity of that city.

Figures

FIG. 1. — Aspects of Xylopia L. vegetative and floral morphology: A, shrub habit of X. humblotiana Baill.; B, habit of X. bemarivensis Diels, showing several nodes with both collateral buds forming shoots, creating pseudo-opposite branches; C, flowers of X. lastelliana Baill., showing inner petals bent outward through the gaps between outer petals (Schatz & Modeste 2877); D, flowers of X.perrieri Diels showing conspicuous red coloration at the bases of the outer petals (Schatz & O’Connor 2964); E, spreading outer petals and erect inner petals of X. kalabenonensis D.M.Johnson, Deroin, & Callmander (Razafitsalama et al. 1041); F, flower of X. richardii Boivin ex Baill., with stigmas emerging in the center amidst the inner petals; G, inflorescences of X. beananensis Cavaco & Keraudren, showing a bud, two flowers with petals, stamens, and stigmas fallen so that the staminal cone is visible, and one open flower (Buerki 14); H, open flower of X. lamarckii Baill.; I, open flower of X. lamarckii with one outer petal and one inner petal removed to show interior of flower, including stamens and stigmas. Photos: A, M. C. Andriamahay; B, F. Rakotoarison; C, D, G. E. Schatz; E, J. Razafitsalama; F, H, I, C. Kaiser-Bunbury; G, M. W. Callmander. A, B, used under the terms of the Creative Commons license for iNaturalist; C-E, G, used under the terms of the Creative Commons license for Tropicos, the botanical information system at the Missouri Botanical Garden.

FIG. 14. — Xylopia bemarivensis Diels, X. ambanjensis, and X. lastelliana Baill. A-K, X. bemarivensis: A, stamen, abaxial view; B, inner petal, adaxial surface; C, outer petal, adaxial surface; D, seed, side view; E, seed, view of micropylar end; F, habit; G, flower, side view; H, bud, side view with one outer petal removed to show inner petals; I, flower with petals, stamens, and stigmas removed to show staminal cone; J, fruit; K, leaf. L-P, X. ambanjensis: L, inflorescence; M, fruit; N, leaves; O, seed, side view; P, seed, view from micropylar end. Q-Y, X. lastelliana: Q, stamen, abaxial view; R, outer staminode, abaxial view; S, T, inner petals adaxial surface; U, outer petal, adaxial surface; V, flower with petals and stamens removed to show stigmas and staminal cone; W, leaf base abaxial surface,

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).

FIG. 29. — Distributions of Xylopia galokothamna, sp. nov., X. ghesquiereana Cavaco & Keraudren, X. humbertii Ghesq. ex Cavaco & Keraudren, X. humblotiana Baill., X. kalabenonensis D.M.Johnson, Deroin, & Callmander, X. lamii Cavaco & Keraudren, X. lastelliana Baill., X. lemurica Diels, X. lokobensis, sp. nov., X. longirostra, sp. nov., X. madagascariensis Cavaco & Keraudren, and X. marojejyana, sp. nov., on Madagascar.