Xylopia perrieri 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 : 74-78

31. Xylopia perrieri Diels

( Figs 1D ; 2B ; 26 )

Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 9: 351 (1925). — Type: Madagascar. Prov. Toamasina [“Madagascaria orientalis”], forêt d’Analamazaotra [c. 18°56’S, 48°25’E], 800 m, XII (fl.), Perrier de la Bâthie 4936 (holo-, P[P030413]!; iso-, P[P030414, P030415]!).

MATERIAL EXAMINED. — Madagascar. Prov. Antananarivo. Centre, Forêt d’Ambohitantely, sur le Tampoketsa d’Ankazobe

[18.161°S, 47.302°E], 1600 m, 12.II.1959 (fl.), Capuron 20165 SF (P[P00169371]); Bongolava ouest de Tsiroanomandidy, VII.1974 (fr.), Morat 4640 (P[P01987127, P01987128, P01987129]).

Prov. Antsiranana. Sava, Andapa, Marojejy, Marojejy National Park, trail between camp 2 and 3, 14°26’S, 49°43’E, 10.XII.2005 (fr.), Ranarivelo & Ravelonarivo RHS 752 ( CAS, K, MO) ; au nord d’Andapa, dans la Réserve naturelle intégrale de Marojejy, aux environs du sommet d’Ambatosoratra, 14°32’S, 49°42’E, 1583 m, 17-24. VI.1994 (fr.), Ravelonarivo 259 ( MO) ; Sous-Préfecture d’Andapa, Commune Rurale d’Andapa, Quartier d’Andasibe, Réserve Spéciale d’Anjanaharibe-Sud, environ de Matsaborimaina, suivant la limite, 14°41’20”S, 49°32’30”E, 867 m, 1.IX.1995 (fr.), Ravelonarivo & Rabesonina 845 ( MO, OWU) ; SW d’Andapa, Réserve Spéciale d’Anjanaharibe-Sud, 14°46’15”S, 49°28’E, 1161-1424 m, 21.III.- 7.IV.1994 (fr.), Ravelonarivo et al. 99 ( MO) ; Sava, Andapa, Doany, Anjialavahely, Antsahabalika situé à 10 km vol d’oiseau au Nord d’Anjialava, 14°13’29”S, 49°23’00”E, 1293 m, 27.II.2006 (fl., fr.), Ravelonarivo et al. 1659 (K n.v., MO) .

Prov. Fianarantsoa. Parc national Ranomafana, near Vohipara, 37.5 km due NE (BRG 46°) of Fianarantsoa, 21°13’33.6”S, 47°22’12.54”E, 1000 m, 4.XI.1997 (fr.), Davis 1058 (K n.v., MO) ; Fianarantsoa , 7 km W of Ranomafana, just S of Namorona River, the Duke University Primate Center study site, 21°16’S, 47°25’E, 1000 m, 5.XII.1988 (fr.), Schatz & Miller 2456 ( BR, K n.v., MO, P[P01987121], WAG) ; vicinity of Ranomafana National Park , near Sahavondronana along road 45, 21°16’S, 47°21’E, 1200 m, 26.VII.1994 (fr.), Turk & Marolahy 592 ( MO) ; Ranomafana National Park , parcelle 3, S of National Road 25 at 7 km W of Ranomafana, Valohoaka trail system, 21°18’S, 47°26’E, 1100-1250 m, 19-21.XI.1992 (fl., fr.), Turk et al. 171 ( MO) .

Prov. Mahajanga. Forêt domaniale à Anjiamazava, versant ouest de la Réserve Spéciale d’Anjanaharibe-Sud, Manandriana, Matsoandakana, Befandriana-nord, Mahajanga, 14°48’S, 49°27.20’E, 961 m, 3-7.II.1997 (fl.), Ravelonarivo et al. 1067 (K n.v., MO, OWU).

Prov. Toamasina. Alaotra-Mangoro Region, Ambatovy, 18°48’16”S, 48°19’54”E, 1133 m, 26.XI.2006 (fr.), Andriantiana 394 (MO); Alaotra Mangoro, Moramanga, Ambohibary , Ampitambe , Antsatsaka forest , 18°48’34”S, 48°16’48”E, 1076 m, 15.V.2009 (immature fr.), Antilahimena et al. 7088 (K n.v., MO) ; Alaotra-Mangoro Region, Moramanga, Ambohibary , Ambadidifanatsy forest , Ambatovy , 18°51’38”S, 48°16’22”E, 966 m, 28.XI.2009 (fr.), Antilahimena et al. 7187 ( MO, OWU) ; 1 km NE of Ankosy, 2 km outside limit of RNI Zahamena , 17°28’28”S, 48°44’12’E, 880 m, 21.I.2000 (fl.), Birkinshaw et al. 636 ( GH, P[P02006076]) ; ouest du village d’Antanandava (P. K. 45 de la route Moramanga-Anosibe ), XI.1968 (st.), Capuron 28450-SF (K) ; East Coast, Betampona Réserve naturelle intégrale, 40 km NW of Toamasina, 17°31’S, 49°07’E, 275- 650 m, 28.IX.1993 (fr.), Lewis & Razafimandimbison 660 ( MO, P[P01987125], WAG) ; Alaotra-Mangoro Region, Ambohilero Forêt Classée , 17°59’33”S, 48°35’51”E, 1200 m, 30.IX.2005 (fr.), Lowry et al. 6624 (K n.v., MO, P[P06774749]) ; Manerinerina sur le Tampoketsa entre l’Ikopa et la Betsiboka, XII.1924 (fl., fr.), Perrier de la Bâthie 16745 (P[P00169370, P01987119]) ; Vohimenakely, Ambodirakauna, Ambodisaina , SE du village Ambodisaina , 17°17’25”S, 48°40’30”E, 850 m, 10.XII.2001 (fr.), Rakotondrafara et al. 95 (A, MO, P[P01986990]) ; Ambatondrazaka, Manakambahiny Est., Sahamalaza , Andrarangabe , 17°43’30”S, 48°46’14”E, 950 m, 4.III.2001 (fl.), Rakotondrajaona et al. 154 ( GH, P[P02006074) ; Toamasina, dans la partie nord-ouest de la Réserve naturelle intégrale de Zahamena, à 1 km au sud-ouest du village d’Antenina , Commune d’Imerimandroso , Ambatondrazaka , 17°30’28”S, 48°46’00”E, 910 m, 15.VIII.1994 (fr.), Randrianjanaka & Zafy 199 (K n.v., MO, OWU) ; Alaotra-Mangoro Region, Moramanga, Andasibe , Andasifahatelo , Ambatoaranana , Corridor Forestier Analamay Mantadia, 18°47’59”S, 48°24’29”S, 1041 m, 25.IV.2012 (immature fr.), Rasoazanany & Tafita 143 ( MO, P[P00903560]) ; Fivondronana, Ambatondrazaka, Firaisana , Antanandava , Ankosy , Ambarikely (3 km à l’Est d’Ankosy et à 2:30 de marche à l’Ouest d’Antenina ), Limite du Parc national de Zahamena , 17°29’S, 48°43’E, 800-850 m, 3.VIII.2003 (fr.), Ratovoson et al. 725 ( MO) ; Alaotra-Mangoro Region, commune rurale d’Andasibe, Fokontany de Menalamba , Marohonkona , Site 2 Ambatovy , 18°50’45”S, 48°18’49”E, 1062 m, 7.VI.2007 (fr.), Ravelonarivo 2514 ( MO) ; Moramanga, Andasibe, Menlamba , entre Analamay and Ambatovy , 18°49’47”S, 48°18’41”E, 1224 m, 23.XI.2005 (fr.), Razafindraibe et al. 126 ( MO) ; Alaotra Mangoro, Moramanga, Lakato , village Manasamena , 19°04’02”S, 48°22’02”E, 1041 m, 19.IX.2007 (fr.), Razanatsima et al. 376 (K n.v., MO) ; Analamazaotra , 1919 (immature fr.), Thouvenot 94 bis (P) .

Prov. Toliara. Reserve No. 11, Andohahela, parcel 1, along trail to Eminyminy, c. 10 km NNE of Hazoambo, 24°45’S, 46°51’E, 600 m, 13.II.1990 (fl.), Schatz & O’Connor 2964 (G, K n.v., MO, P[P01954118], PRE, WAG).

Province unknown. Central Madagascar, without definite locality, Baron 1632 (BM, K n.v.).

DISTRIBUTION, ECOLOGY AND CONSERVATION STATUS. — Xylopia perrieri occurs in scattered localities from northern to southeastern Madagascar in humid forests from 600 to 1600 m ( Fig. 30 ). Associates include species of Weinmannia ( Cunoniaceae ), Monimiaceae , and Lauraceae in the canopy, as well as species of Psychotria, Oncostemum, and Cyathea in the understory. Specimens with flowers have been collected in January, February, March, October, and November, and with fruits in all months of the year. Xylopia perrieri was calculated to have an EOO of 50 587 km 2 and an AOO of 120 km 2. Known from 30 localities, this species has the third largest EOO and AOO sizes of any Madagascar species. Although not of highest conservation concern, this species grows in mid-elevation forests that are vulnerable to development pressures and in-country assessment is needed. We have given it a preliminary conservation assessment of Data Deficient ( Table 2 ).

LOCAL NAMES. — Fontsimavo (Thouvenot 94 bis), hazoambo (multiple collections), ramiavona ( Turk & Marolavy 592, Turk et al. 171).

DESCRIPTION

Tree up to 20(-25) m tall; d.b.h. up to 30(-40) cm.

Twigs pubescent, the hairs 0.1-0.7 mm long, eventually glabrate; nodes with one or often two axillary branches.

Leaves with larger blades 2.4-5.9 cm long, 1.5-2.1 cm wide, chartaceous to subcoriaceous, slightly discolorous, green to purplish brown adaxially, light brown abaxially, elliptic to lanceolate, occasionally ovate, apex obtuse, emarginate, or occasionally with a blunt acumen 2.5-5 mm long, base cuneate to broadly cuneate and decurrent on petiole, margin flat, not revolute, glabrous except for the pubescent midrib adaxially, uniformly appressedpubescent abaxially; midrib with age darkened adaxially, secondary veins weakly brochidodromous, 9-14 per side, diverging at 50-80° from midrib, these and higher-order veins forming a conspicuous reticulum on both surfaces; petiole 1.3-4.5 mm long, shallowly canaliculate or flattened, smooth, pubescent.

Inflorescences axillary, 1(-3)-flowered, pedunculate or pedicels arising separately from leaf axil, appressed-pubescent; peduncle c. 0.8 mm long; pedicels 2 per peduncle, 3.6-6.8 (8-11.3) mm long, 0.8-1.0 mm thick; bracts 2(-4), attached near midpoint of pedicel, caducous or the distal bract sometimes persistent, 1.5-1.8 mm long, ovate; buds lanceolate, apex obtuse.

Sepals slightly spreading at anthesis, ¹⁄5-1/2-connate, 1.8-2.9 mm long, 2.6-2.7 mm wide, coriaceous, semicircular to triangular, apex broadly acute to apiculate, sparsely pubescent abaxially.

Petals yellowish-white, red at the base in vivo; outer petals more or less erect at anthesis, 10-16.4 mm long, 2.6-3.7 mm wide at base, 2.0- 2.7 mm wide at midpoint, coriaceous, lanceolate, flat, apex obtuse, pubescent except for glabrous basal concavity adaxially, pubescent abaxially; inner petals more or less erect at anthesis, 8-13.7 mm long, 2.0- 2.5 mm wide at base, 1.3-1.7 mm wide at midpoint, coriaceous, lanceolate, keeled on apical ¼-1/3 adaxially, keeled abaxially, apex acute, base concave with undifferentiated margin, puberulent except for glabrous base adaxially, puberulent abaxially.

Stamens c. 180; fertile stamens 1.2-1.4 mm long, narrowly oblong to clavate, anther connective apex 0.2-0.3 mm long, shield-shaped, overhanging anther thecae, papillate, anthers 10-11-locellate, filament c. 0.4 mm long; outer staminodes 1.0- 1.2 mm long, clavate, apex obtuse; inner staminodes 1.1-1.2 mm long, clavate, apex rounded; staminal cone 1.7- 2.1 mm in diameter, 0.7-1.0 mm high, partially to completely concealing the ovaries, rim irregularly laciniate.

Carpels 7-11;ovaries 1.2-1.4 mm long, oblong, pubescent, stigmas loosely connivent, 3.7-4.5 mm long, filiform, glabrous to pilose.

Fruit of up to 8 monocarps borne on a pedicel 6-12 mm long, 2.5-6 mm thick, glabrate; torus 4.9-20 mm in diameter, 3.8- 10 mm high, irregularly depressed-globose; monocarps with yellowish red to reddish green exterior and rose-colored to orange-red endocarp in vivo, 1.9-4.2 cm long, 1.3-2.1 cm wide, 1.4-1.9 cm thick, oblong to pyriform, often weakly torulose, apex rounded, base sessile or contracted into a stipe 3-6 mm long, 4-6 mm thick, verrucose, longitudinally or obliquely wrinkled, usually shiny, glabrate; pericarp 0.4-1.8 mm thick.

Seeds up to 7 per monocarp, in two rows, sometimes interdigitated to form a single row, more or less perpendicular to long axis, 12.2-14.4 mm long, 8.5-10.9 mm wide, 5.6-6.9 mm thick, ovoid, semicircular to wedge-shaped in cross-section, light brown to reddish brown, smooth, dull or slightly shiny, perichalazal ring not evident; sarcotesta waxy, light green in vivo; aril absent.

NOTES

Xylopia perrieri may be recognized by its small leaves, which are usually densely appressed-pubescent abaxially, obtuse flower buds, lanceolate petals strongly marked with red at the base, and oblong shiny monocarps. The buds and flowers most resemble those of X. humbertii , but the leaves of X. humbertii are glabrous. Xylopia perrieri also resembles X. buxifolia , but the latter species has acute rather than obtuse flower buds, linear rather than lanceolate petals, and leaves usually widest at or distal to the midpoint. The seeds of X. perrieri lack the donut-shaped thickening around the micropyle usually present on the seeds of X. buxifolia . Collections of X. perrieri come from elevations of 600-1600 m, while those of X. buxifolia are from elevations below 500 meters, rarely reaching 900 meters. Xylopia perrieri is sympatric with X. lemurica in some localities, for example Ranomafana National Park, but lacks the abruptly acuminate leaves, longer and more persistent hairs of the leaf midrib and margins, and needle-like inner petals found in that species.

Xylopia perrieri is remarkably uniform in morphology throughout its range. The specimen Schatz & O’Connor 2964 from Toliara Province varies from other collections in its longer pedicels, some reaching over 11 mm in length, and in bearing 3-4 bracts rather than the usual 2 on these longer pedicels. The collection is identical to others of the species, however, in its other floral features, including the bicolored outer petals described by most collectors ( Fig. 1D ).

Several Xylopia specimens from elevations between 400 and 1000 meters from Toamasina Province resemble X. perrieri in leaf form but have monocarps that are not wrinkled or torulose but instead are finely verrucose and have a longitudinal ridge down one side of the monocarp: Capuron 28155-SF (K n.v., P), Est, à l’Est de Marozevo (Beforona), 23.I.1968 (fr.); Poncy et al. 1565 (K, MO, P[P00373087]), Maroantsetra, Tampolo, Péninsule Masoala, entre Ambodiforaha et Laotrozo, 15°42’17”S, 49°58’43”E, 400-450 m, 2.XI.2001 (buds, fr.); Ravelonarivo et al. 4148 (MO); Atsinanana Region, Vohimanana, Dist.Brickaville, Com. Maroseranana, Fkt. Ambatolampy, 18°23’36”S, 48°48’38”E, 942 m, 23.I.2012 (fr.). Two specimens from the same area and with the same fruit morphology also have immature flower buds, which are more acute than is typical for X. perrieri: Randrianjanaka & Arnaud 9 (G, K, MO, P[P01987123]),Toamasina, Réserve naturelle intégrale 3, Zahamena, au NW du village Andratanantsoitra, 17°43’S, 48°59’E, 700 m, 9-14.XII.1993 (fl. buds, fr.); Rasoanindriana 385 (OWU), District Toamasina II, Commune Rurale de Sahambala, Fokontany de Sahavongo, village de Sahavongo, forêt d’Andasibe, 18°01’39”S 49°07’16”E, 648 m, 24.XI.2017 (buds, fr.). Also problematic is the specimen Razafitsalama & Torze 1149 (K, MO), Ambilobe, Beramanja, Anketrabe, forêt de Kalabenono, sur la chaîne Galoko, 7 km au SE d’Anketrabe, 13°39’09”S, 48°40’35”E, 795 m, 25.XI.2006 (fr.). The leaf blades of the specimen fit those of X. perrieri in both shape and indument although they are somewhat large for the species, and the monocarps are torulose and have the same shiny wrinkled surface. However, the specimen did not cluster with the accession of X. perrieri included in our molecular analysis ( Stull et al. 2017). Additional field observations are needed to determine the status of all these specimens.

The type and other specimens of this species at P collected by Perrier de la Bâthie were annotated in 1934 by Ghesquière with the name Xylopia lastelliana var. perrieri (Diels) Ghesq. As far as we can determine, this combination was never published.

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. 2. — Fruits and seeds of Madagascar Xylopia L. species: A, fruits of X. retusa, sp. nov., showing at left a dehiscent monocarp with one seed covered by an orange sarcotesta (Rabenantoandro et al. 1249); B, fruit of X. perrieri Diels, one monocarp dehiscing to reveal barely visible seed with light green sarcotesta (Antilahimena 7187); C, dehiscing monocarps of a Xylopia sp., possibly X. carinata, sp. nov., with orange sarcotesta of seeds visible; D, fruit of X. bemarivensis Diels, with one dehiscing monocarp and seed with orange sarcotesta visible (Ranirison PR 808); E, immature fruit of X. flexuosa Diels, showing the elongate monocarp characteristic of species in sect. Xylopia (Rakotovao 4687); F, fruit of X. humblotiana Baill., with one dehisced monocarp and black seeds with white arils visible (Rakotovao 4886); G, fruits of X. sclerophylla, sp. nov.; monocarp at lower right is probably recently dehisced because green sarcotesta of seed is fully intact (Nusbaumer LN 943); H, dehisced monocarp of X. sclerophylla, sp. nov., at a slightly later stage, showing the sarcotesta collapsing and losing its color (Nusbaumer LN 943). Photos: A, P. P. Lowry II; B, P. Antilahimena; C, J. Tan; D, P. Ranirison (© Conservatoire et Jardin botaniques de la Ville de Genève); E, F, C. Rakotovao; G, H, by L. Nusbaumer (© Conservatoire et Jardin botaniques de la Ville de Genève). A, B, D, E, F, used under the terms of the Creative Commons license for Tropicos,the botanical information system at the Missouri Botanical Garden; C, used under the terms of the the Creative Commons license for iNaturalist.

FIG. 26. — Xylopia perrieri Diels: A, abaxial surface of leaf base, showing indument; B, fruit; C, inner petal, adaxial view; D, outer petal, adaxial view; E, stamen, abaxial view; F, habit; G, calyx and gynoecium, lateral view;H, seed, view from micropylar end; I, seed, lateral view.A, C-G, from Ravelonarivo et al. 1067 (OWU); B, H, I, from Schatz & Miller 2456 (MO). Scale bars: A, B, F, 1 cm; C-E, G, 1 mm; H, I, 5 mm.

FIG. 30. — Distributions of Xylopia perrieri Diels, X. ravelonarivoi, sp. nov., X. retusa, sp. nov., X. sclerophylla, sp. nov., X. sericolampra Diels and X. sahafariensis Cavaco & Keraudren on Madagascar.