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