Xylopia lemurica Diels, 1925

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 : 70-72

27. Xylopia lemurica Diels

( Fig. 17L-N )

Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 9: 350 (1925). — Type: Madagascar. Prov. Toamasina [“Madagascaria centralis”], Analamazaotra , in silvis, 800 m, II.1912 (fl., fr.), Perrier de la Bâthie 4975 (lecto-, here designated, P[P030406]!; isolecto-, B [B100153146]!).

Xylopia pseudolemurica Cavaco & Keraudren , Bulletin de la Société botanique de France 103: 276 (1956). — Type: Madagascar. Prov. Antsiranana, Massif de l’Ambohitsitondroina de Mahalevona , nord de la presqu’île de Masoala, vers 400 m d’alt., 28.XI.1953 (fl.), Capuron 8674 (lecto-, here designated, P [ P030416 ]!; iso-, OWU!, P[P030417, P030418]).

Xylopia microphylla Cavaco & Keraudren , Bulletin de la Société botanique de France 103: 277 (1956). — Type: Madagascar. Prov. Toamasina [“ Domaine de l’Est ”], massif du Beanjada, N de la presqu’île Masoala, vers 1000 m d’alt., 28.XII.1953 (fl.), Capuron 8815 (holo-, P [ P030407 ]!; iso-, P[P030408, P030409]!).

MATERIAL EXAMINED. — Madagascar. Prov. Antsiranana. SW of Andapa, near SW edge of Anjanaharibé-Sud Reserve, 14°48’15”S, 49°26’45”E, 1000-1100 m, 6.VIII.1997 (fr.), McPherson 17265 ( MO); Marombihy , Mandena , Parc national de Marojejy, le long d’un affluent de la rivière Manantenina, 11 km du village Manantenina, 14°26.2’S, 49°44.5’E, 1200 m, 1.XI.1996 (fl., fr.), Messmer et al. NM 312 (G, MO) ; Préfecture d’Antalaha , Sous-Préfecture d’Andapa , Commune rurale de Bealampona, Quartier de Befingotra , Sud Ouest d’Andapa , Réserve spéciale d’Anjanaharibe-Sud , suivant la route nationale d’Andapa-Bealanana, environ d’Ampiferantany, 14°47’45”S, 49°27’45”E, 1020 m, 3.VII.1995 (immature fr.), Ravelonarivo & Remainty 829 ( MO, OWU, P[P01954125]) .

Prov. Fianarantsoa. Ranomafana National Park, parcelle 3, S of National Road 25 at 7 km W of Ranomafana, Talatakely trail system, 21°15’30”S, 47°25’00”E, 950-1150 m, 8.XII.1992 (fl.), Turk & Radriamanatena 202 ( MO); Ranomafana National Park , parcelle 3, S of National Road 25 at 7 km W of Ranomafana, Talatakely trail system, 21°15’30”S, 47°25’00”E, 950-1150 m, 26.I.1993 (fl., fr.), Turk & Solo 262 ( MO, OWU) .

Prov. Mahajanga. Sous-Préfecture de Befandriana-Nord, commune rurale de Matsoandakana, Quartier de la Belalona, Sud-Ouest d’Andapa, Réserve spéciale Anjanaharibe-Sud, village d’Anjiamazava, suivant la route nationale d’Andapa-Bealanana, piste vers le Nord approchant le sommet de Bevitsika, 14°42’S, 49°27’E, 1100 m, 14.XII.1994 (fl.), Ravelonarivo & Rabesonina 549 (OWU).

Prov. Toamasina. Fivondronana: Moramanga, Commune: Andasibe, Fokontany: Menalamba, Ambatovy Forest, 18°50’13”S, 48°19’19”E, 1130 m, 17.I.2005 (fl., fr.), Antilahimena et al. 3208 (K n.v., MO, P[P02006079]); Fivondronana : Moramanga , Commune : Andasibe , Fokontany : Berano, Ambatovy Forest, 18°48’29”S, 48°18’45”E, 1120 m, 26.I.2005 (fl.), Antilahimena et al. 3243 ( MO, OWU) ; Fivondronana : Moramanga , Commune : Andasibe , Fokontany : Berano, Ambatovy Forest, 18°49’14”S, 48°20’07”E, 1110 m, 3.II.2005 (fl., fr.), Antilahimena et al. 3265A ( MO, P[P06901344]) ; Alaotra-Mangoro Region , Moramanga , Commune Ambohibary, Fokontany Ampitambe, 18°51’02” S, 48°17’24”E, 1080 m, 9.II.2007 (fl.), Antilahimena & Edmond 5245 (K n.v., MO, P[P06901346]) ; Phelps Dodge project site, c. 15 aire-km NE of Moramanga, c. 11 km E of Antanambao, between Ambatovy-South and Analamay-East , Torotorofotsy River, Berano, 18°50’32”S, 48°19’55”E, 1000 m, 18.II.1997 (fr.), Andriatsiferana et al. 2184 ( MO, P[P02006077], WAG) ; Alaotra Mangoro , Moramanga , Ambohibary , Ampitambe, Ambatovy, 18°49’S, 48°18’E, s. d. (immature fr.), Andriatsiferana et al. 2489 ( MO) ; ouest du village d’Antanandava (P. K. 45 de la route Moramanga-Anosibe, XI.1968 (fr.), Capuron 28412-SF (K) ; Phelps Dodge project site, c. 15 aire-km NE of Moramanga, c. 11 km E of Antanambao, between Ambatovy-South and Analamay-East , Torotorofotsy River, Berano, 18°50’32”S, 48°19’55”E, 1000 m, 20.II.1997 (fl.), Rakotomalaza et al. 1145 ( MO) ; Vatomandry , Ambalabe , Ambinaninandro II, direction sud ouest de Toby Foara, 19°09’36”S, 48°34’44”E, 636 m, 20.I.2006 (fl.), Razanatsima & Honoré 82 (K n.v., MO) ; Fiv. Moramanga , Ambatovy, Sahavaharina Forest, 18°51’32”S, 48°20’44”E, 975 m, 3.II.2006 (fl.), Rogers & Antilahimena 1019 ( OWU) .

DISTRIBUTION, ECOLOGY AND CONSERVATION STATUS. — Xylopia lemurica occurs in eastern Madagascar, from Antsiranana Province south to Fianarantsoa Province in humid forests along the eastern slope at elevations of 400-1150 meters, most commonly from over 1000 m ( Fig. 29 ). This species is one of several, also including X. beananensis , X. danguyella , and X. perrieri , that frequent lower montane forests. The labels of the two Turk collections from Ranomafana describe the habitat as humid montane forest characterized by trees of Weinmannia spp., Monimiaceae (Tam - bourissa, Decarydendron, Ephippiandra ), Lauraceae (primarily species of Ocotea and Cryptocarya), with Psychotria spp., Oncostemum spp., and Cyathea spp. abundant in the understory, a similar list of associates as given for Xylopia perrieri at the same site. The MO sheet of Turk & Solo 262 shows three spiny galls, which appear to have developed from flowers. Similar “flower galls” occur in some African species, such as X. mwasumbii ( Johnson et al. 1999) . Specimens with flowers have been collected from November to February, and with fruits in July, August, and from November to February. With an EOO value of 38 756 km 2 and an AOO value of 64 km 2, the species has a relatively large range but is seemingly uncommon throughout, and is given a preliminary conservation assessment of Data Deficient.

LOCAL NAMES. — Fotsimavo (Perrier de la Bâthie 4975), hazoambo (six collections), ramiavona ( Turk & Radriamanatena 202, Turk & Solo 262).

DESCRIPTION

Tree up to 17 m tall; d.b.h. up to 27 cm.

Twigs finely appressed-pubescent, the hairs 0.1-0.8 mm long, becoming glabrate; twigs often branching with expansion of new growth, nodes with one or frequently two axillary branches.

Leaves with larger blades 3.5-8.3 cm long, 1.4-2.8 cm wide, chartaceous to subcoriaceous, discolorous, purplish brown adaxially, brown abaxially, elliptic to lanceolate, apex acuminate, the acumen 3-11 mm long and often bent to the side when pressed, base cuneate and short-decurrent on petiole, margin flat, not revolute, glabrous except for erect hairs on midrib adaxially, sparsely appressed-pubescent with the hairs longest, densest, and most persistent on the midrib and margin abaxially; midrib light brown toward base adaxially, secondary veins irregularly brochidodromous, 11-15 per side, diverging at 50-60° from midrib, these and higherorder veins indistinct to slightly raised adaxially, raised and forming a fine reticulum abaxially; petiole 1.5-6 mm long, shallowly canaliculate, longitudinally wrinkled, sparsely pubescent to glabrate.

Inflorescences axillary, 1-2-flowered, pedunculate or pedicels arising separately from leaf axil, densely fine-pubescent; peduncle 0.8-2 mm long; pedicels 2 per peduncle, 3.1-9.3 mm long, 0.5-0.8 mm thick; bracts 2-3, 2 attached near midpoint and sometimes a third just above the pedicel base, caducous or rarely persistent, 1-2 mm long, broadly ovate to semicircular; buds linear-subulate to linear-lanceolate, apex obtuse, occasionally falciform.

Sepals spreading at anthesis, ¹⁄5-1/2-connate, 1.5-2.7 mm long, 1.8-2.6 mm wide, coriaceous, triangular, apex acute, pubescent abaxially.

Petals yellow, red to maroon at base on both surfaces in vivo; outer petals erect or slightly curved outward at anthesis, 10.2- 20.3 mm long, 1.9-3.3 mm wide at base, 0.7-1.4 mm wide at midpoint, fleshy, linear, flat or slightly concave adaxially, keeled except at the base abaxially, apex obtuse, puberulent adaxially, sericeous abaxially; inner petals perhaps slightly curved outward at anthesis, 8.3-18 cm long, 1.6-2.4 mm wide at base, 0.6-1.0 mm wide at midpoint, fleshy, linear, keeled except at the base adaxially, keeled to the base abaxially, apex acute to obtuse, base deeply concave with undifferentiated margin but somewhat bilobed at widest point, puberulent only along keel adaxially, puberulent abaxially.

Stamens 120-160; fertile stamens 0.8-1.2 mm long, oblong, anther connective apex 0.2-0.3 mm long, shield-shaped, overhanging anther thecae, papillate, anthers 6-8-locellate, filament 0.2-0.5 mm long; outer staminodes 1.0- 1.1 mm long, oblong, apex rounded or truncate; inner staminodes 0.7-0.8 mm long, oblong or broadly clavate, apex rounded, truncate, or umbonate; staminal cone 1.1-1.9 mm in diameter, 0.6-1.1 mm high, concealing all but the apices of the ovaries, rim irregularly laciniate.

Carpels 6-9; ovaries 1-1.2 mm long, narrowly oblong, densely pubescent, stigmas loosely connivent, 2.2-2.7 mm long, filiform, sparsely pubescent.

Fruit of up to 7 monocarps borne on a pedicel 6-15 mm long, 2.1-6 mm thick, glabrate; torus 7-12 mm in diameter, 4-8 mm high, depressed-globose; monocarps with a pink to red exterior and pinkish red endocarp in vivo, 1.5-2.9 cm long, 1.1-1.5 cm wide, 1.2-1.4 cm thick, ellipsoid, oblong, or obovoid, slightly torulose, apex obtuse to rounded, base sessile, slightly narrowed, smooth, longitudinally wrinkled, dull, with a longitudinal adaxial ridge, glabrate; pericarp 0.7-1.5 mm thick.

Seeds up to 6 per monocarp, in two rows or sometimes a single irregular row, perpendicular to long axis, 8.5-10.3 mm long, 5.4-7.4 mm wide, 4.5-5.8 mm thick, ellipsoid to oblong-ellipsoid, slightly thickened around the micropyle, semicircular to wedge-shaped in cross-section, brown to light brown, smooth, dull, perichalazal ring forming a slight ridge on antiraphal side; sarcotesta yellow-green in vivo; aril absent.

NOTES

Xylopia lemurica may be distinguished by the relatively small elliptic to lanceolate and usually abruptly acuminate leaves, the needle-like inner petals, and the relatively small blunt sessile and torulose monocarps often marked by an adaxial ridge. The hairs along the leaf margins and abaxial midrib tend to be both longer and more persistent than on the rest of the leaf. Xylopia perrieri , which occurs in many of the same lower montane sites (both were described from the same type locality, Analamazaotra), may be distinguished by the obtuse leaves with denser persistent abaxial pubescence of uniform length, the broader petals, and larger and shinier monocarps with larger seeds. In the linear sparsely pubescent petals and shape of the monocarps X. lemurica more closely resembles X. buxifolia , but the sharp leaf acumen of X. lemurica will distinguish the two.

Cavaco & Keraudren (1956) distinguished Xylopia pseudolemurica from X. lemurica on the basis of the color of the indument on the young branches and petioles (“chamois-clair”), by the larger fawn-velutinous calyx, and the longer petals. Xylopia microphylla was distinguished from X. lemurica , in the same publication, by its leaves densely pubescent with silvery hairs, and from X. pseudolemurica by its smaller leaves, and the calyx smaller and lacking the fawn-colored hairs. With additional collections it is now possible to see that all of these characters represent extremes of variation. Two collections from Ranomafana National Park illustrate the situation. The specimen Turk & Radriamanatena 202 bears flowers with sepals c. 2.7 mm long but outer petals only c. 11.5 mm in length, thus not in keeping with the combination of characteristics used by Cavaco and Keraudren to separate X. lemurica and X. pseudolemurica , and has a relatively coarse sericeous indument on the leaves and twigs of silvery hairs 0.7-0.8 mm long, which seems to be quickly lost. The specimen Turk & Solo 262, in contrast, has flowers with sepals c. 1.9 mm long and outer petals 16.5-19 mm long, and finer pubescence (hairs only 0.2-0.3 mm long) on the twigs and leaves. The type specimen of X. lemurica does, in fact, have relatively short hairs and short petals compared to the type specimen of X. pseudolemurica .

Neither the P sheet nor the B sheet of Perrier de la B â thie 4975, the type collection of Xylopia lemurica Diels , is identified as the holotype. Both bear drawings of floral details, presumably by Diels. The P sheet is the more ample material, and shows a greater study with respect to the detail drawings attached, and is chosen as the lectotype.

The type information given in the protologue for Xylopia pseudolemurica is “Domaine de l’Est: nord de Tamatave, Service Forestier 7888 (type).” A second collection is also listed: “Domaine de l’Est: […] massif de l’Ambohitsitondroina de Mahalevona, Capuron 8674.” The type locality and number appear to have been an error, as this information has been crossed out on the holotype sheet and the collection details for the Capuron 8674 specimen substituted. A note has also been glued to the sheet, stamped with “Service des Eaux et Forets, Madagascar ”, stating that the sheet cannot be 7888 SF. The specimen Capuron 8674 is therefore understood to be the type of this name, but the sheet P030416 is designated here as the lectotype to avoid future confusion.

A chromosome number of 2n=16 was reported for this species by Morawetz & Le Thomas (1988) but the voucher collections, Le Thomas 15 and Le Thomas 20, have not been traced. Given the low elevation provenance of the collections, however, it is likely that the collections do not represent this species.

Figures

FIG. 17. — Xylopia buxifolia Baill. and X. lemurica Diels. A-K, X. buxifolia; A, leaves; B, fruit; C, seed, side view; D, seed, view of micropylar end; E, outer staminode; F, fertile stamen; G, inflorescence; H, outer petal, adaxial surface; I, inner petal, adaxial surface; J, flower with petals and most of stamens removed to show staminal cone and stigmas; K, bud, side view; L-N, X. lemurica; L, flower bud, side view; M, open flower, side view; N, habit. A-K, from Schatz & Carlson 2861; A, WAG; E, F, H-K, MO,sheet 1 of 2;B-D, G, MO sheet 2 of 2;L-N, from Perrier de la Bâthie 4975 (P). Scale bars:A, B, G,K-N,1 cm;C, D,5 mm;E,F,1 mm;H,I, 3 mm;J, 2 mm.

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.