Xylopia perrieri Diels 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 Diels Diels 1925 [1038,1272,1715,1742] Magnoliopsida Annonaceae Xylopia Plantae Magnoliales 75 74 Tracheophyta species perrieri   ( Figs 1D; 2B; 26)    Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem9: 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 Bâthie4936(holo-, P[P030413]!; iso-, P[P030414, P030415]!).  MATERIAL EXAMINED. —   Madagascar. Prov. Antananarivo. Centre, Forêt d’Ambohitantely, sur le Tampoketsa d’Ankazobe  FIG. 25. —  Xylopia longirostra, sp. nov., and  X. marojejyana, sp. nov. A -E,  X. longirostra, sp. nov.: A, habit; B, close-up of rolled apex of developing leaf; C, immature flower bud, side view; D, monocarp, side view; E, seed, side view. F -H,  X. marojejyana, sp. nov.: F, habit; G, monocarp, adaxial view; H, petiole and twig, side view. A, D, E, from Lowry et al. 6064(OWU); B, from Razakamalala 266(NY); C, from Andrianjafy et al. 120(MO); F -H, based on Miller & Randrianasolo 4423(MO). Scale bars: A, D-G, 1 cm; B, H, 2 mm; C, 1 mm. [ 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 Réservenaturelle intégrale de Marojejy, aux environs du sommet d’Ambatosoratra, 14°32’S, 49°42’E,  1583 m,  17-24. VI.1994(fr.),  Ravelonarivo259( 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& Rabesonina845( 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.),  Ravelonarivoet al. 99( MO);  Sava, Andapa, Doany, Anjialavahely, Antsahabalikasitué à 10 kmvol d’oiseau au Nord d’Anjialava, 14°13’29”S, 49°23’00”E,  1293 m,  27.II.2006(fl., fr.),  Ravelonarivoet al. 1659(K n.v., MO).   Prov. Fianarantsoa.Parc national Ranomafana, near Vohipara, 37.5 kmdue 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 kmW of Ranomafana, just S of Namorona River, the Duke University Primate Centerstudy site, 21°16’S, 47°25’E,  1000 m,  5.XII.1988(fr.),  Schatz& Miller2456( BR, K n.v., MO, P[P01987121], WAG); vicinity of Ranomafana National Park, near Sahavondronanaalong road 45, 21°16’S, 47°21’E,  1200 m,  26.VII.1994(fr.),  Turk& Marolahy592( MO);  Ranomafana National Park, parcelle 3, S of National Road25 at  7 kmW of Ranomafana, Valohoakatrail system, 21°18’S, 47°26’E,  1100-1250 m,  19-21.XI.1992(fl., fr.),  Turket 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.),  Antilahimenaet 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.),  Antilahimenaet al. 7187( MO, OWU);   1 kmNE of Ankosy, 2 kmoutside limit of RNI Zahamena, 17°28’28”S, 48°44’12’E,  880 m,  21.I.2000(fl.),  Birkinshawet al. 636( GH, P[P02006076]); ouest du village d’Antanandava(P. K. 45 de la route Moramanga-Anosibe),  XI.1968(st.),  Capuron28450-SF(K);  East Coast, Betampona Réservenaturelle intégrale,  40 kmNW of Toamasina, 17°31’S, 49°07’E,  275- 650 m,  28.IX.1993(fr.),  Lewis& Razafimandimbison660( MO, P[P01987125], WAG);  Alaotra-Mangoro Region, Ambohilero Forêt Classée, 17°59’33”S, 48°35’51”E,  1200 m,  30.IX.2005(fr.),  Lowryet al. 6624(K n.v., MO, P[P06774749]);  Manerinerina sur le Tampoketsaentre l’Ikopaet la Betsiboka,  XII.1924(fl., fr.),  Perrier de la Bâthie16745(P[P00169370, P01987119]);  Vohimenakely, Ambodirakauna, Ambodisaina, SE du village Ambodisaina, 17°17’25”S, 48°40’30”E,  850 m,  10.XII.2001(fr.),  Rakotondrafaraet 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.),  Rakotondrajaonaet al. 154( GH, P[P02006074);  Toamasina, dans la partie nord-ouest de la Réservenaturelle intégrale de Zahamena, à 1 kmau 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& Zafy199(K n.v., MO, OWU);  Alaotra-Mangoro Region, Moramanga, Andasibe, Andasifahatelo, Ambatoaranana, Corridor Forestier AnalamayMantadia, 18°47’59”S, 48°24’29”S,  1041 m,  25.IV.2012(immature fr.),  Rasoazanany& Tafita143( MO, P[P00903560]);  Fivondronana, Ambatondrazaka, Firaisana, Antanandava, Ankosy, Ambarikely( 3 kmà l’Est d’Ankosyet à 2:30 de marche à l’Ouest d’Antenina), Limite du Parcnational de Zahamena, 17°29’S, 48°43’E,  800-850 m,  3.VIII.2003(fr.),  Ratovosonet al. 725( MO);  Alaotra-Mangoro Region, commune rurale d’Andasibe, Fokontany de Menalamba, Marohonkona, Site2 Ambatovy, 18°50’45”S, 48°18’49”E,  1062 m,  7.VI.2007(fr.),  Ravelonarivo2514( MO);  Moramanga, Andasibe, Menlamba, entre Analamayand Ambatovy, 18°49’47”S, 48°18’41”E,  1224 m,  23.XI.2005(fr.),  Razafindraibeet al. 126( MO);  Alaotra Mangoro, Moramanga, Lakato, village Manasamena, 19°04’02”S, 48°22’02”E,  1041 m,  19.IX.2007(fr.),  Razanatsimaet al. 376(K n.v., MO);  Analamazaotra, 1919 (immature fr.),  Thouvenot94 bis(P).   Prov. Toliara.Reserve No. 11, Andohahela, parcel 1, along trail to Eminyminy, c. 10 kmNNE 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 perrierioccurs in scattered localities from northern to southeastern Madagascarin humid forests from 600 to 1600 m( Fig. 30). Associates include species of Weinmannia( Cunoniaceae), Monimiaceae, and Lauraceaein the canopy, as well as species of Psychotria, Oncostemum, and Cyatheain 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 perrieriwas calculated to have an EOO of 50 587 km 2and an AOO of 120 km 2. Known from 30 localities, this species has the third largest EOO and AOO sizes of any Madagascarspecies. 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,  Turket al. 171).  DESCRIPTION  Treeup to 20(-25) m tall; d.b.h. up to 30(-40) cm.  Twigspubescent, the hairs 0.1-0.7 mmlong, eventually glabrate; nodes with one or often two axillary branches.  Leaveswith larger blades 2.4-5.9 cmlong, 1.5-2.1 cmwide, 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 mmlong, 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 mmlong, shallowly canaliculate or flattened, smooth, pubescent.  Inflorescencesaxillary, 1(-3)-flowered, pedunculate or pedicels arising separately from leaf axil, appressed-pubescent; peduncle c. 0.8 mmlong; 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 mmlong, ovate; buds lanceolate, apex obtuse.  Sepalsslightly spreading at anthesis, ¹⁄5-1/2-connate, 1.8-2.9 mmlong, 2.6-2.7 mmwide, coriaceous, semicircular to triangular, apex broadly acute to apiculate, sparsely pubescent abaxially.  Petalsyellowish-white, red at the base in vivo; outer petals more or less erect at anthesis, 10-16.4 mmlong, 2.6-3.7 mmwide at base, 2.0- 2.7 mmwide 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 mmlong, 2.0- 2.5 mmwide at base, 1.3-1.7 mmwide 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 mmlong, narrowly oblong to clavate, anther connective apex 0.2-0.3 mmlong, shield-shaped, overhanging anther thecae, papillate, anthers 10-11-locellate, filament c. 0.4 mmlong; outer staminodes 1.0- 1.2 mmlong, clavate, apex obtuse; inner staminodes 1.1-1.2 mmlong, clavate, apex rounded; staminal cone 1.7- 2.1 mmin diameter, 0.7-1.0 mm high, partially to completely concealing the ovaries, rim irregularly laciniate.  Carpels7-11;ovaries 1.2-1.4 mmlong, oblong, pubescent, stigmas loosely connivent, 3.7-4.5 mmlong, filiform, glabrous to pilose.  Fruitof up to 8 monocarps borne on a pedicel 6-12 mmlong, 2.5-6 mmthick, glabrate; torus 4.9-20 mmin diameter, 3.8- 10 mmhigh, irregularly depressed-globose; monocarps with yellowish red to reddish green exterior and rose-colored to orange-red endocarp in vivo, 1.9-4.2 cmlong, 1.3-2.1 cmwide, 1.4-1.9 cmthick, oblong to pyriform, often weakly torulose, apex rounded, base sessile or contracted into a stipe 3-6 mmlong, 4-6 mmthick, verrucose, longitudinally or obliquely wrinkled, usually shiny, glabrate; pericarp 0.4-1.8 mmthick.  Seedsup 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 mmlong, 8.5-10.9 mmwide, 5.6-6.9 mmthick, 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 perrierimay 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. humbertiiare glabrous.  Xylopia perrierialso 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. perrierilack the donut-shaped thickening around the micropyle usually present on the seeds of  X. buxifolia. Collections of  X. perriericome from elevations of 600-1600 m, while those of  X. buxifoliaare from elevations below 500 meters, rarely reaching 900 meters.  Xylopia perrieriis sympatric with  X. lemuricain 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.  FIG. 26. —  Xylopia perrieriDiels: 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.   Xylopia perrieriis remarkably uniform in morphology throughout its range. The specimen Schatz & O’Connor 2964from Toliara Province varies from other collections in its longer pedicels, some reaching over 11 mmin 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  Xylopiaspecimens from elevations between 400 and 1000 meters from Toamasina Province resemble  X. perrieriin leaf form buthave 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 kmau 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. perrieriin 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. perrieriincluded in our molecular analysis ( Stull et al.2017). Additional field observations are needed to determine the status of all these specimens. The typeand other specimens of this species at P collected by Perrier de la Bâthie were annotated in 1934 by Ghesquière with the name  Xylopia lastellianavar. perrieri(Diels) Ghesq. Asfar as we can determine, this combination was never published. 2632071716 Perrier de la Bathie Madagascar 800 -18.933332 d'Analamazaotra 1273 48.416668 75 74 1 Toamasina holotype 2632071484 75 74 2632071576 75 74 2632071671 76 75 2632071738 1994-06-17 1994-06-24 1994-06-17 MO d'Andapa & Reserve & Marojejy & d'Ambatosoratra 1583 -14.533334 Ravelonarivo 1287 49.7 76 75 1 2632071328 1995-09-01 MO, OWU Sous-Prefecture d'Andapa & Commune Rurale d'Andapa & Quartier d'Andasibe & Reserve Speciale d'Anjanaharibe-Sud & Matsaborimaina & Ravelonarivo & Rabesonina 867 -14.688889 21 49.541668 76 75 1 2632071439 1994-03-21 1994-04-07 1994-03-21 MO SW d'Andapa & Reserve Speciale d'Anjanaharibe-Sud 1293 -14.770833 Ravelonarivo 894 49.466667 76 75 1 2632071453 2006-02-27 MO Sava & Andapa & Doany & Anjialavahely & Antsahabalika & Nord d'Anjialava 1293 -14.224722 Ravelonarivo 21 49.383335 76 75 1 2632071648 76 75 2632071672 1988-12-05 BR, MO, WAG Schatz & Miller 1000 -21.266666 Fianarantsoa 1264 47.416668 Duke University Primate Center 76 75 1 2632071568 1994-07-26 MO Marolahy Turks and Caicos Islands 1200 -21.266666 Ranomafana National Park 1264 47.35 Sahavondronana 76 75 1 2632071510 1992-11-19 1992-11-21 1992-11-19 MO Turks and Caicos Islands 1175 -21.3 Ranomafana National Park 1264 47.433334 Valohoaka 76 75 1 2632071719 76 75 2632071589 76 75 2632071683 2009-05-15 MO Antilahimena Alaotra Mangoro 1076 -18.809444 Ambohibary 21 48.28 Moramanga 76 75 1 2632071383 2009-11-28 MO, OWU Antilahimena Alaotra-Mangoro Region 966 -18.860556 Ambohibary 21 48.272778 Moramanga 76 75 1 2632071318 2000-01-21 GH Birkinshaw 880 -17.474445 Zahamena 76 75 1 2632071505 1968-11 Capuron Moramanga-Anosibe d'Antanandava 76 75 1 2632071350 1993-09-28 MO, WAG Lewis & Razafimandimbison 463 -17.516666 Betampona Reserve 1278 49.116665 East 76 75 1 Pwani 2632071612 2005-09-30 MO Lowry 1200 -17.9925 Ambohilero Foret Classee 21 48.5975 Alaotra-Mangoro Region 76 75 1 Pwani 2632071520 1924-12 l'Ikopa & la Betsiboka & Perrier de la Bathie Manerinerina sur le Tampoketsa 76 75 1 Pwani 2632071468 2001-12-10 MO Rakotondrafara Vohimenakely 850 -17.290277 Ambodisaina 21 48.675003 Ambodirakauna 76 75 1 Pwani 2632071381 2001-03-04 GH Rakotondrajaona Ambatondrazaka 950 77 76 -17.725 Sahamalaza 21 48.770554 Manakambahiny Est. 76 75 1 Pwani 2632071479 1994-08-15 MO, OWU Randrianjanaka & Zafy Reserve 910 -17.507778 d'Antenina 21 48.766666 Zahamena 77 76 1 Toamasina 2632071403 2012-04-25 MO Rasoazanany & Tafita Alaotra-Mangoro Region 1041 -48.40806 Andasibe Moramanga 77 76 1 Toamasina 2632071613 2003-08-03 MO Ratovoson Fivondronana 825 -17.483334 Firaisana 1278 48.716667 Ambatondrazaka 77 76 1 Toamasina 2632071525 2007-06-07 MO Ravelonarivo Alaotra-Mangoro Region 1062 -18.845835 Fokontany de Menalamba 21 48.31361 d'Andasibe 77 76 1 Toamasina 2632071524 2005-11-23 MO Razafindraibe Moramanga 1224 -18.829723 Menlamba 21 48.31139 Andasibe 77 76 1 Toamasina 2632071698 2007-09-19 MO Razanatsima Alaotra Mangoro 1041 -19.067223 Lakato 21 48.36722 Moramanga 77 76 1 Toamasina 2632071597 [132,698,772,796] Thouvenot Analamazaotra 77 76 1 Toamasina 2632071313 77 76 2632071624 77 76