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21. Annonaceae

番荔枝科 fan li zhi ke

Authors: Bingtao Li, Michael G. Gilbert & George Schatz

Trees, shrubs, or climbers, wood and leaves often aromatic; indument of simple or less often (Uvaria, Annona) stellate hairs. Leaves alternate, normally distichous. Stipules absent. Petiole usually short; leaf blade simple, venation pinnate, margin entire. Inflorescences terminal, axillary, leaf-opposed, or extra-axillary [rarely on often underground suckerlike shoots]. Flowers usually bisexual, less often unisexual, solitary, in fascicles, glomerules, panicles, or cymes, sometimes on older wood, usually bracteate and/or bracteolate. Sepals hypogynous, [2 or]3, imbricate or valvate, persistent or deciduous, rarely enlarging and enclosing fruit, free or basally connate. Petals hypogynous, 3-6(-12), most often in 2 whorls of 3 or in 1 whorl of 3 or 4[or 6], imbricate or valvate, sometimes outer whorl valvate and inner slightly imbricate. Stamens hypogynous, usually many, rarely few, spirally imbricate, in several series; filaments very short and thick; anther locules 2, contiguous or separate, rarely transversely locular, adnate to connective, extrorse or lateral, very rarely introrse, opening by a longitudinal slit; connectives often apically enlarged, usually ± truncate, often overtopping anther locules, rarely elongated or not produced. Carpels few to many, rarely solitary, free or less often connate into a 1-locular ovary with parietal placentas; ovules 1 or 2 inserted at base of carpel or 1 to several in 1 or 2 ranks along ventral suture, anatropous; styles short, thick, free or rarely connate; stigmas capitate to oblong, sometimes sulcate or 2-lobed. Fruit usually apocarpous with 1 to many free monocarps, these sometimes moniliform (constricted between seeds when more than 1-seeded), often fleshy, indehiscent, rarely dehiscent (Anaxagorea, Xylopia), and often with base extended into stipe, rarely on slender carpophore (Disepalum), less often syncarpous with carpels completely connate and seeds irregularly arranged and sometimes embedded in fleshy pulp. Seeds 1 to many per monocarp or many per syncarp, often arillate; endosperm copious, ruminate; embryo minute, near hilum.

About 129 genera and over 2,300 species: tropics and subtropics, especially in Old World; 24 genera (one endemic, three introduced) and 120 species (41 endemic, 11 introduced) in China.

The flowers often open before the petals have fully expanded making it easy to underestimate floral dimensions.

One of the major studies of the Annonaceae of tropical Asia was by Suzanne Jovet-Ast (1914-2006), who published an account of the Indochinese species under her maiden name, Ast (Fl. Indo-Chine, Suppl. 1: 62-121. 1938). Unfortunately, she failed to provide Latin descriptions for her many new taxa, none of the names of which was therefore validly published. Her new combinations were validly published, however, and she did subsequently validate a few species of Goniothalamus under her maiden name; but she did not validate most of her new species until after she married and took up the name "Jovet-Ast." These later names form the major part of all her publications, so we have chosen to cite her as "Jovet-Ast ["Ast"]" for those taxa validated under her maiden name.

Tsiang Ying & Li Ping-t’ao. 1979. Annonaceae. In: Tsiang Ying & Li Ping-t’ao, eds., Fl. Reipubl. Popularis Sin. 30(2): 10-175.


1 Fruit syncarpous, with completely united carpels and seeds embedded in pulp (usually cultivated as fruit trees)   (2)
+ Fruit apocarpous, with carpels forming free, often stipitate, monocarps   (3)
       
2 (1) Outer petals usually free, abaxially flat.   23 Annona
+ Outer petals connate at base, abaxially spurred or winged.   24 Rollinia
       
3 (1) Sepals or inner petals and sometimes also outer petals clearly imbricate; stellate hairs often present   (4)
+ Sepals and inner and outer petal whorls all valvate or rarely very narrowly imbricate at tip only (only 1 whorl of petals present in Dasymaschalon); simple hairs usually present, less often plant glabrous   (6)
       
4 (3) Trees or self-supporting shrubs; stellate hairs absent; sepals imbricate; petals valvate or very minutely imbricate; ovules 1 per carpel.   15 Trivalvaria
+ Climbing shrubs; stellate hairs present; sepals valvate; inner petals and sometimes also outer petals imbricate; ovules 2 or more per carpel   (5)
       
5 (4) Petals spreading at anthesis; torus depressed.   2 Uvaria
+ Petals incurved at anthesis, not opening widely; torus flattened, margin prominent.   3 Cyathostemma
       
6 (3) Climbing shrubs   (7)
+ Trees or erect shrubs   (10)
       
7 (6) Peduncles and fruiting pedicels hooklike.   20 Artabotrys
+ Peduncles and fruiting pedicels not hooklike   (8)
       
8 (7) Monocarps ellipsoid, moniliform when more than 1-seeded; petals basally constricted to form enclosed floral chamber around stamens and carpels.   6 Desmos
+ Monocarps globose to cylindric, never moniliform; petals not constricted   (9)
       
9 (8) Monocarp stipes 5.3-7.5 cm; outer petals usually much larger than inner petals; stamen connectives apically truncate (flowers of F. hainanensis unknown).   21 Friesodielsia
+ Monocarp stipes 0.3-4 cm; outer petals only slightly larger than inner petals; stamen connectives apically acuminate to obtuse.   22 Fissistigma
       
10 (6) Petals (2 or)3, in 1 whorl.   7 Dasymaschalon
+ Petals 6, in 2 whorls   (11)
       
11 (10) Inner petals basally clawed or stipitate, laminas vaulted and connate into an apical cap at least initially   (12)
+ Inner petals not basally clawed, spreading or connivent and apically 3-edged   (16)
       
12 (11) Outer petals 2-7 mm, usually smaller than inner petals and similar to sepals (inner and outer petals subequal in Orophea hainanensis and O. yunnanensis)   (13)
+ Outer petals more than 12 mm, equaling or longer than inner petals, conspicuously different from sepals   (14)
       
13 (12) Inner petals with a long basal claw, usually apically lozenge-shaped and cohering by their margins into a cap over stamens and carpels, sometimes with their tips conspicuously recurved; pedicel 0.2-1.2 cm.   4 Orophea
+ Inner petals with a short basal claw, shallowly saccate, at first cohering by their margins but soon free; pedicel usually 1.4-10 cm (0.5-0.7 cm in M. glochidioides).   5 Miliusa
       
14 (12) Stamens linear-oblong; inner petals each with a short claw or stipe.   8 Goniothalamus
+ Stamens cuneate; inner petals each with a long claw or stipe   (15)
       
15 (14) Outer petals 1.6-2.5(-3.4) cm, longer than inner petals.   9 Mitrephora
+ Outer petals 1-1.5 cm, shorter than inner petals.   10 Pseuduvaria
       
16 (11) Stamen connectives apically acuminate   (17)
+ Stamen connectives apically truncate, rounded, or broadly 3-angular   (19)
       
17 (16) Petals 5-8 cm, linear to linear-lanceolate.   19 Cananga
+ Petals 0.7-1 cm, ovate-triangular to ovate-oblong, usually saccate and incurved at base   (18)
       
18 (17) Monocarps dehiscent, stipe clavate; ovules 2 per carpel.   1 Anaxagorea
+ Monocarps indehiscent, stipe not clavate; ovules 5-22 per carpel.   18 Alphonsea
       
19 (16) Ovules 1 or 2(or 3) per carpel, basally or nearly basally inserted   (20)
+ Ovules many per carpel, ventrally inserted   (22)
       
20 (19) Inner petals concave and incurved to cover stamens; fruiting carpels free but usually connivent.   17 Popowia
+ Inner petals spreading; fruiting carpels free and not connivent   (21)
       
21 (20) Monocarp bases extended into stipes, not articulate at base.   14 Polyalthia
+ Monocarps on slender carpophores, articulate between apex of carpophore and base of monocarp.   16 Disepalum
       
22 (19) Flower buds lanceolate, apically long beaked and 3-angled; sepals connate at base or beyond into a cup; petals elongated, concave at base, narrowly convex below middle; anther locules transversely septate.   11 Xylopia
+ Flower buds ovoid, apically rounded; sepals connate at base but not into a cup; petals flat or cucullate; anther locules not transversely septate   (23)
       
23 (22) Petals flat; stamen connectives apically rounded; stigmas subcapitate, not constricted at base; monocarps almost sessile (Chinese species).   12 Meiogyne
+ Petals cucullate; stamen connectives apically truncate to subtruncate; stigmas ovoid, constricted at base; monocarps long stipitate.   13 Chieniodendron

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