Neotropical Annonaceae

Paul J.M. Maas

National Herbarium, Wageningen University Branch, the Netherlands. 

Description

Trees, shrubs, or very rarely lianas, bark very tough and fibrous and easily torning off in long strips, often with a spicy aroma when cut. Indument of vegetative, floral, and fruiting parts composed of simple, stellate, or lepidote hairs, or lacking. Leaves simple, alternate, distichous, very rarely spirally arranged (Tetrameranthus R.E.Fr.),  pinnately veined. Flowers axillary or internodal (often leaf-opposed), rarely terminal, single or in rhipidiate inflorescences, rarely in panicles, bracts present, sometimes lacking (a.o. Cymbopetalum Benth.); sepals (2-)3(-4), free or variously connate; petals 6, rarely 3 (e.g. some species of Annona L.) or 8 (Stenanona p.p. and Tetrameranthus), mostly in two 3-merous whorls, free, rarely connate; stamens many, spirally arranged, filaments mostly very short, anthers dithecic, extrorsely and longitudinally dehiscent, sometimes transversely septate (a.o. Xylopia L.) often with an apical, shield-like prolongation of the connective; staminodes rarely present; carpels many, rarely few, with a sessile stigma or apex more or less narrowed into a style, placentation basal or lateral, rarely apical, ovules 1 to many. Fruit mostly apocarpous, the carpels (monocarps) usually stipitate, fleshy, indehiscent, sometimes dehiscent (e.g. Anaxagorea St.Hil. and Xylopia), less frequenly fruit pseudosyncarpous (e.g. Duguetia A.St.-Hil.) or syncarpous (e.g. Annona). Seeds 1-many per monocarp, often rather large, with ruminate endosperm, sometimes arillate.

Notes on delimitation

  • The family of Annonaceae is placed in the order of Magnoliales, together with 2 families which also occur in the Neotropics, namely Myristicaceae and Magnoliaceae.

Distribution in the Neotropics

  • All over the Neotropics, in the North even to the USA and S Canada (Asimina Adans.), particularly common in the lowlands.

Distinguishing characters (always present)

  • Vegetatively easily recognizable by its wood showing concentric bands of parenchyma and by its bark, which is very tough and fibrous and can be torn off in long strips.
  • Leaves distichous.
  • Flowers are (in 95% of the cases) 3-merous, consisting of 3 sepals and 6 (much larger) petals.
  • Stamens and free carpels spirally arranged on the receptacle.
  • Endosperm ruminate.

Other important characters

  • Stamens generally many, mostly with an apical shield-like prolongation of the connective.
  • Fruit mostly apocarpous.

Key differences from similar families

Annonaceae can hardly be confused with the two other Neotropical families belonging to the order of Magnoliales, namely Magnoliaceae and Myristicaceae by the above mentioned characters of wood and bark:

Number of genera

35 genera:

  • Anaxagorea (21 spp.)
  • Annona (incl. Raimondia Saff. and Rollinia A.St.-Hil.) (200 spp.)
  • Asimina (10 spp.)
  • Bocagea A.St.-Hil. (3 spp.)
  • Bocageopsis R.E.Fr. (4 spp.)
  • Cardiopetalum Schltdl. (3 spp.)
  • Cremastosperma R.E.Fr. (29 spp.)
  • Cymbopetalum (27 spp.)
  • Desmopsis Saff. (17 spp.)
  • Diclinanona Diels (3 spp.)
  • Duckeanthus R.E.Fr. (1 sp.)
  • Duguetia (90 spp.)
  • Ephedranthus S.Moore (10 spp.)
  • Froesiodendron R.E.Fr. (3 spp.)
  • Fusaea Saff. (2 spp.)
  • Guatteria Ruiz & Pav. (incl. Guatteriella R.E.Fr., Guatteriopsis R.E.Fr., and Heteropetalum Benth. ) (175 spp.)
  • Hornschuchia Nees (10 spp.)
  • Klarobelia Chatrou (9 spp.)
  • Malmea R.E.Fr. (6 spp.)
  • Mosannona Chatrou (14 spp.)
  • Onychopetalum R.E.Fr. (2 spp.)
  • Oxandra A.Rich. (25 spp.)
  • Porcelia Ruiz & Pav. (6 spp.)
  • Pseudephedranthus Aristeg. (1 sp.)
  • Pseudomalmea Chatrou (3 spp.)
  • Pseudoxandra R.E.Fr. (22 spp.)
  • Ruizodendron R.E.Fr. (1 sp.)
  • Sapranthus Seem. (10 spp.)
  • Stenanona Standl. (13 spp.)
  • Tetrameranthus (6 spp.)
  • Tridimeris Baill. (1 sp.)
  • Trigynaea Schltdl. (8 spp.)
  • Unonopsis R.E.Fr. (49 spp.)
  • Xylopia (40 spp.)

Useful tips for generic identification

 

Follow link to illustrated key to Neotropical Annonaceae

 

http://www.kew.org/science/tropamerica/neotropikey/families/annonaceae_illustrated_key.pdf

 

Key to genera of Neotropical Annonaceae

1.  Leaves spirally arranged; flowers 4(-6)-merous; indument composed of stellate hairs. The Amazon region and French Guiana in the North and the Pacific coast of Colombia in the West … Tetrameranthus
1.  Leaves arranged in two rows along the stem (distichous); flowers 3-merous (very rarely 2-merous); indument of various types (simple, stellate, or scale-like hairs) … 2.

2.  Indumentum composed of stellate or scale-like hairs (well visible with a hand lens); fruit pseudo-syncarpous and carpels strongly coherent, but only partly connate, the lower carpels sterile and often forming a basal collar. From Costa Rica in the North to Bolivia and Paraguay in the South … Duguetia
2.  Indumentum composed of simple hairs or lacking (hairs sometimes stellate in Annona, microscopic stellate to furcate hairs occur in Anaxagorea); fruit either apocarpous or syncarpous, but not as above … 3.

3.  Flowers and fruit axillary (sometimes leaf-opposed in Anaxagorea brevipes) or sometimes arising from the trunk (cauliflorous); fruit apocarpous … 4.
3.  Flowers and fruit terminal, leaf-opposed, or supra-axillary, rarely arising from the trunk; fruit apocarpous or syncarpous … 22.

4.  Upper leaf surface with impressed midrib … 5.
4.  Upper leaf surface with raised midrib … 14.

5.  Petals often thick and fleshy (2-10 mm thick) and covered with microscopic, brownish hairs; monocarps club-shaped, explosively dehiscent, 2-seeded. All over the Neotropics, except for the West Indian islands … Anaxagorea
5.  Petals generally much thinner and covered with much larger hairs or glabrous; monocarps never club-shaped, generally globose to ellipsoid, indehiscent, or non-explosively dehiscent, 1-several-seeded … 6.

6.  Pedicels with a supra-basal articulation (a few milimetres above the base of the pedicel). All over the Neotropics … Guatteria
6.  Pedicels with a basal articulation … 7.

7.  Sepals 2; petals 4; carpels 1(-2). Mexico … Tridimeris
7.  Sepals 3; petals 6; carpels generally numerous … 8.

8.  Monocarps dehiscent, 1-several-seeded; seeds distinctly arillate; young twigs mostly lenticellate; leaves generally small and narrow. All over the Neotropics …  Xylopia
8.  Monocarps indehiscent, 1-several-seeded; aril absent or indistinct; young twigs not lenticellate; leaves mostly much larger and broader … 9.

9.  Monocarps several-seeded … 10.
9.  Monocarps 1-seeded … 11.

10.  Petals often maroon, 10-80 x 10-50 mm, distinctly veined; inner petals much shorter than the outer ones; monocarps 1-12. SE USA to S Canada … Asimina 10.  Petals mostly white, 7-23 x 1-3 mm, not veined; petals subequal or inner petals longer than the outer ones; monocarps 1-2. Tropical South America, but mainly Amazonian … Diclinanona

11.  Upper leaf surface with distinctly impressed venation; monocarps long-stipitate (stipes 5-35 mm long); petals 8-12 mm long. Tropical South America, but mainly in NE Brazil … Ephedranthus
11.  Upper leaf surface with flat, not or slightly impressed venation; monocarps short- to long-stipitate; petals 4-70 mm long … 12.

12.  Pedicels with 3-6 bracts; stipes of monocarps less than 8 mm long; petals 4-8 mm long; all flowers bisexual. All over the Neotropics … Oxandra
12.  Pedicels with 1-2 bracts; stipes of monocarps more than 10 mm long, generally much longer; petals 7-70 mm long; flowers androdioecious (male and bisexual flowers present) … 13.

13.  Pedicels with 1 bract above the articulation; petals 8-21 mm long, spreading and leaving the floral center uncovered; raphe of seeds raised, straight. Western South America … Pseudomalmea
13.  Pedicels without a bract above the articulation; petals 10-30 mm long, concave, covering the floral center; raphe of seeds impressed, straight to slightly sinuous. Tropical South America up to Cost Rica in the North … Klarobelia

14.  Petals often fleshy (2-10 mm thick) and covered with microscopic, brownish hairs; innermost stamens staminodial; monocarps club-shaped, explosively dehiscent, 2-seeded. All over the Neotropics, except for the West Indian islands … Anaxagorea
14.  Petals generally much thinner and covered with much larger hairs or glabrous; all stamens fertile or outermost stamens staminodial (Fusaea); monocarps globose to ellipsoid, indehiscent or non-explosively dehiscent, 1-several-seeded … 15.

15.  Leaves with distinct marginal vein, almost touching the margin (except in P. espirito-santensis); petals 4-15 mm long. Tropical South America …  Pseudoxandra
15.  Leaves without a marginal vein (except in Oxandra p.p., but then much further removed from the margin); petals 4-35 mm long … 16.

16.  Lower leaf surface glaucous; monocarps transversely ellipsoid, 1-seeded; petals 25-35 mm long. Western part of South America … Ruizodendron
16.  Lower leaf surface not glaucous; monocarps ellipsoid to globose, 1-several-seeded; petals 4-20 mm long … 17.

17.  Monocarps 1-2, sessile, 2-4-seeded; petals 4-8.5 mm long, with a small, incurved, tail-like, apical appendage. Tropical South America … Onychopetalum
17.  Monocarps generally many more (up to 30), mostly distinctly stipitate, often 1-seeded; petals 2.5-20 mm long, without an apical appendage … 18.

18.  Leaves asymmetrical; petals 2.5-4.5 mm long; monocarps 1-seeded (except in B. pleiosperma), shortly stipitate. Tropical South America … Bocageopsis
18.  Leaves symmetrical; petals mostly > 5 mm long; monocarps 1-several-seeded, mostly distinctly stipitate … 19.

19.  Upper leaf surface with a distinctly grooved midrib. From Costa Rica in the North to Bolivia in the South, eastwards up to the Guianas, and S of the Amazon River … Cremastosperma
19.  Upper leaf surface with a non-grooved midrib … 20.

20.  Leaves with distinctly raised veins on both surfaces; petals 10-15 mm long; seeds 25-30 mm long. Amazonian SW Venezuela and adjacent Brazil ... Pseudephedranthus
20.  Leaves with veins on the upper surface (except for the raised midrib) not or indistinctly raised; petals 4-10 mm long; seeds 8-20 mm long ... 21.

21.  Inflorescences often 1-flowered, the pedicel densely beset with 3-6 bracts; apical prolongation of connective tongue-shaped; monocarps 1-seeded. All over the Neotropics … Oxandra
21.  Inflorescences often branched, if 1-flowered, the pedicel not densely beset with bracts; apical prolongation of connective discoid; monocarps 1-several-seeded. From Mexico (Oaxaca) in the North to Bolivia, N Paraguay, and S Brazil in the South ... Unonopsis

22.  Bracts absent … 23.
22.  Bracts present … 29.

23.  Flowers pendent on long (40-60 mm) pedicels; inner petals boat-shaped. All over the Neotropics, except for the West Indian islands … Cymbopetalum
23.  Flowers non-pendent and pedicels much shorter; inner petals not boat-shaped … 24.

24.  Leaves asymmetrical; monocarps 20-90 x 30-40 mm, wall 2.5-4 mm thick. From Costa Rica in the North through Western South America to Bolivia, and SE Brazil … Porcelia
24.  Leaves symmetrical; monocarps smaller, wall thinner … 25.

25.  Petals basally connate … 26.
25.  Petals free … 27.

26.  Petals basally connate into a 2-4 mm long tube; monocarps dehiscent, falciform, yellow or orange; aril fleshy, 2-lobed. Tropical South America … Cardiopetalum
26.  Petals free; monocarps indehiscent, fusiform to linear, green; aril fibrous. Western South America … Froesiodendron

27.  Leaves tri-plinerved at the base; monocarps 3-9; flower buds globose; stamens c. 40. Tropical South America, lacking in the central Amazon basin … Trigynaea 27.  Leaves not tri-plinerved at the base; monocarps 1-3; flower buds globose or conical; stamens < 20. E and SE Brazil … 28.

28.  Flower buds conical; petals linear, recurved to patent; calyx cup-shaped, without distinct lobes; flowers solitary or in many-flowered inflorescences; inflorescences often flagellate and produced from the main stem. E and SE Brazil … Hornschuchia
28.  Flower buds globose; petals ovate to elliptic, erect; calyx distinctly 3-lobed; flowers solitary. E and SE Brazil … Bocagea

29.  Upper leaf surface with raised midrib … 30.
29.  Upper leaf surface with impressed midrib … 31.

30.  Fruit apocarpous, the monocarps distinctly stipitate; seeds pitted; sepals < 5 mm long; base of inner petals with food bodies. From Mexico in the North to Peru in the South … Mosannona
30.  Fruit pseudo-syncarpous, the carpels strongly coherent; seeds smooth; sepals > 9 mm long; petals without food bodies. Amazonian Brazil (Upper Rio Negro) … Duckeanthus

31.  Flowers winged; fruit syncarpous (very rarely apocarpous). All over the Neotropics. … Annona p.p.
31.  Flowers not winged; fruit apocarpous or syncarpous … 32.

32.  Outermost stamens staminodial; fruit syncarpous and provided with an almost woody basal collar. Tropical South America ... Fusaea
32.  All stamens fertile; fruit apocarpous or syncarpous, but then without a basal collar … 33.

33.  Fruit syncarpous, each carpel 1-seeded; petals often thick and fleshy. All over the Neotropics … Annona
33.  Fruit apocarpous, each monocarp 1-several-seeded; petals thin … 34.

34.  Pedicels often provided with 1 leafy bract. Mexico and Central America and adjacent N Colombia … Desmopsis
34.  Pedicels without leafy bracts … 35.

35.  Monocarps 1-seeded; petals yellow to cream, margins ciliate. From Panama in the North to Peru in the South, also 1 species in SE Brazil (Bahia) … Malmea
35.  Monocarps 1-several-seeded; petals red, purple, brown, or rarely cream, not ciliate … 36.

36.  Petals distinctly veined, length-width ratio 2-5; flowers often with a fetid scent; monocarps 13-100 mm long; seeds with lamellate rumination. Mexico and Central America … Sapranthus
36.  Petals not distinctly veined, length-width ratio 5-20; flowers without a fetid scent; monocarps 10-30 mm long; seeds with spiniform rumination. Mexico, Central America to the Pacific coast of Colombia … Stenanona

Notable genera and distinguishing features

  • Anaxagorea: Neotropical genus which also occurs in tropical Asia. Unique in the family by its explosively dehiscent (2-seeded) monocarps; the seeds can in this way been dispersed up to many meters (7 m) from the mother tree!
  • Annona: Genus also occurring in Africa. Recent molecular and morphological research by H. Rainer (in prep.) has revealed that the genera Rollinia and Raimondia should be merged into Annona. The genus Annona is characterized by having syncarpous fruits (except for some species of the former genus Rollinia). It occurs all over the Neotropics, also in the dry cerrado vegetations of Brazil, where most other genera of Annonaceae are lacking.
  • Cymbopetalum: As the name Cymbo (boat) and petalum (petal) already suggests this genus is characterized by having boat-shaped inner petals (with the margins mostly strongly rolled inwards). Furthermore its flowers are mostly placed on long, pendent, ebracteate pedicels.
  • Duguetia: Recognizable by its indument of stellate to lepidote hairs. The fruits are peudosyncarpous (carpels connate in various degrees). A few species of Duguetia, which were formerly placed in the genus Pachypodanthium Engl. & Diels, occur in W tropical Africa.
  • Fusaea: This small genus of only two species is quite remarkable by relatively large staminodes (placed between the inner petals and stamens) and by a syncarpous fruit.
  • Guatteria: The largest genus in the Neotropics, which is currently under revision in Brazil (A. Lobao), Utrecht (R.H.J. Erkens) and Wageningen (P.J.M. Maas & L.Y.Th. Westra). It is best recognizable by having axillary flowers with a pedicel which has a suprabasal articulation, whereas all other Neotropical genera have a pedicel with basal articulation.
    It is a nightmare to study this genus in the herbarium, as mostly only "open flower buds" are present. In Guatteria the flower starts as a very young closed bud, then they open and the flowers stay open and green for a long period of several weeks. After that the petals start to lengthen, they change from green to yellow, become erect and enclose with their base a so-called pollination chamber. Only then the pollination takes place. In herbarium material there are almost always only open flower buds present, which means that the measurements of "petals" are often incorrect.
  • Hornschuchia: A genus restricted to E and SE Brazil. Various species have their flowers arranged in long flagellate inflorescences (a.o. H. bryotrophe Nees).
  • Oxandra: Good characters to distinguish this genus are found in the shortly pedicellate flowers, in which the pedicels are densely covered with various bracts. In many species the apical prolongation of the connective is acute (Oxy (acute) and andros(male)).
  • Porcelia: Genus with very large, many-seeded monocarps, reaching a length of up to 13 cm.
  • Tetrameranthus: Unique within the Neotropical genera of Annonaceae by having spirally arranged leaves and 4-merous flowers.
  • Xylopia: The only genus in the family occurring in the 3 continents, America, Africa, and Asia. The genus is characterized by axillary flowers, in which the outer petals are generally much larger than the inner ones. The monocarps are dehiscent, mostly several-seeded, and the seeds are provided with a distinct aril.

 

Status

  • All genera are native in the Neotropics.
  • Several species are cultivated all over the Neotropics because of their edible fruits: some species of Annona, a.o. A. muricata L., A. cherimola Miller, A. mucosa Jacq. (the former Rollinia mucosa), and A. reticulata L.
  • Young stems of Annonaceae are often used as fishing rods.

General notes

In a few species of the family the flowers are arranged in a flagellate inflorescence. This occurs in Duguetia, Hornschuchia, Stenanona, and in 1 species of Guatteria.

Important literature

Chatrou, L.W. 1998. Changing genera. Systematic studies in Neotropical and West African Annonaceae. pp.1-224. PhD thesis. Utrecht.

Couvreur, T.L.P., Maas, P.J.M., Meinke, S., Johnson, D.M. & Kessler, P.J.A. 2012. Keys to the genera of Annonaceae. Bot. J. Linn. Soc. 169(1): 74-83.

Erkens, R.H.J. 2007. From morphological nightmare to molecular conundrum. Phylogenetic, evolutionary and taxonomic studies on Guatteria (Annonacae).  pp.1-276. PhD thesis. Utrecht.

Johnson, D.M. & N.A. Murray. 1995. Synopsis of the tribe Bocageeae (Annonaceae) with revisions of Cardiopetalum, Froesiodendron, Trigynaea, Bocagea, and Hornschuchia. Brittonia 47: 248-319.

Kessler, P.J.A. 1993. Annonaceae, pp 93-129. In: K. Kubitzki (ed.), The families and genera of vascular plants vol 2. Springer-Verlag, Berlin.

Maas, P.J.M. & L.Y.Th. Westra 1984/85. Studies in Annonaceae II. A monograph of the genus Anaxagorea A. St.-Hil. Bot. Jahrb. Syst. 105: 73-134. 1984; Bot Jahrb. Syst. 105: 145-204. 1985.

Maas, P.J.M. & L.Y.Th. Westra.1992. Rollinia (Annonaceae). Flora Neotropica Monograph 57: 1-188.

Maas, P.J.M. & L.Y.Th. Westra. 2003. Revision of the Neotropical genus Pseudoxandra. Blumea 48: 201-259.

Maas, P.J.M. , L.Y.Th. Westra and L.W. Chatrou. 2003. Duguetia (Annonaceae). Flora Neotropica Monograph 88: 1-274.

Maas, P.J.M., L.Y.Th. Westra & M. Vermeer. 2007. Revision of the Neotropical genera Bocageopsis, Onychoptalum, and Unonopsis (Annonaceae). Blumea 52: 413-554.

Murray, N.A. 1993. Revision of Cymbopetalum and Porcelia (Annonaceae). Syst. Bot. Monogr. 40: 1-121.

Oliveira, J. & M.F. Sales. 1999. Estudo taxonomico dos generos Ephedranthus S. Moore e Pseudephedranthus Aristeg. - Annonaceae. Bol. Mus. Paraense Emilio Goeldi, n.s., Bot. 15(2): 117-166.

Pirie, M.D.  2005. Cremastosperma (and other evolutionary digressions). Molecular phylogenetic, biogeographic, and taxonomic studies in Neotropical Annonacae. pp.1-256. PhD thesis. Utrecht.

Westra, L.Y. Th. 1985. Studies in Annonaceae IV. A taxonomic revision of Tetrameranthus R.E. Fries. Proc. Kon. Ned. Akad. Wetensch. C. 88: 449-482.

How to cite

Maas, P.J.M. (2009). Neotropical Annonaceae. In: Milliken, W., Klitgεrd, B. & Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://www.kew.org/science/tropamerica/neotropikey/families/Annonaceae.htm.

Click images to enlarge


Anaxagorea dolichocarpa © L. Campbell, RBG, Kew.



Anaxagorea dolichocarpa © L. Campbell, RBG, Kew.



Fruit of Annona cherimola © M. Ackermann, Freie Universitδt Berlin.



Annona cherimola, flowering branch © M. Weigend, Freie Universitδt, Berlin.



Fruit of Annona hypoglauca © D. Sasaki, Programa Flora Cristalino.



Annona macrocalyx © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Annona montana © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Annona muricata © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Annona scandens © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Fruits of Cymbopetalum brasiliense © D. Sasaki, Programa Flora Cristalino.



Cymbopetalum brasiliense © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Cymbopetalum longipes © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Cymbopetalum sp. © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Duguetia confusa © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Duguetia flagellaris © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Fruits of Duguetia marcgraviana © W. Milliken, RBG, Kew.



Developing fruits of Duguetia riparia © D. Sasaki, Programa Flora do Cristalino.



Duguetia spixiana © D. Sasaki, Programa Flora do Cristalino.



Duguetia spixiana © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Flower of Fusaea longifolia © D. Sasaki, Programa Flora do Cristalino.



Fusaea longifolia © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Guatteria allenii © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Guatteria burchellii © A.M. Amorim.



Guatteria pudica © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Guatteria pudica © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Foliage and sapwood of Oxandra xylopioides © D. Sasaki, Programa Flora do Cristalino.



Unonopsis bullata © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Unonopsis duckei © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Unonopsis guatterioides © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Unonopsis guatterioides © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Xylopia aromatica © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Xylopia aromatica © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.



Xylopia peruviana © P.J.M. Maas, National Herbarium, Wageningen Branch, the Netherlands.