The families of flowering plants

L. Watson and M. J. Dallwitz

Annonaceae Juss.
Including Hornschuchiaceae J.G. Agardh, Monodoraceae J.G. Agardh
Habit and leaf form. Trees,

or shrubs, or lianas; bearing essential oils; resinous, or

not resinous. Self supporting, or climbing; when climbing, scrambling, or stem
twiners, or petiole twiners. Leaves evergreen; alternate;distichous; nonsheathing; gland-dotted, or not gland-dotted; aromatic, or without marked odour;
simple. Lamina entire; pinnately veined; cross-venulate. Leaves exstipulate.
Lamina margins entire. Domatia occurring in the family (recorded in 3 genera);
manifested as pockets (usually), or hair tufts (in 1/12 species).
Leaf anatomy. The

leaf lamina dorsiventral. Abaxial epidermis papillose, or not

papillose. Stomata paracytic. Adaxial hypodermis absent. Lamina with secretory
cavities, or without secretory cavities. Secretory cavities containing oil, or
containing mucilage, or containing resin. The mesophyll usually with spherical
etherial oil cells (?); containing mucilage cells, or not containing mucilage cells;
with sclerenchymatous idioblasts, or without sclerenchymatous idioblasts. Main
veins vertically transcurrent. Minor leaf veins without phloem transfer cells
(Annona).
Axial (stem, wood) anatomy. Young

stems with solid internodes. Pith with

diaphragms (commonly, comprising stone cells), or without diaphragms;
heterogeneous (with groups of stone cells even in the absence of complete septa).
Secretory cavities present; with resin. Cork cambium present; initially
superficial. Nodes unilacunar (with three traces), or bilacunar (according to
Lammers et al 1986). Primary vascular tissues in a cylinder, without separate
bundles, or comprising a ring of bundles; collateral. Internal phloem absent.
Cortical bundles absent. Medullary bundles absent. Secondary thickening
developing from a conventional cambial ring. Primary medullary rays wide, or
mixed wide and narrow, or narrow. The axial xylem with vessels (but these rather
few).
The wood variously ring porous to diffuse porous. The vessels very small;
solitary, radially paired, in radial multiples, and clustered. The vessel end-walls
horizontal; simple. The vessels without vestured pits; with spiral thickening
(rarely, reported in Asimena), or without spiral thickening. The axial xylem with
fibre tracheids. The fibres without spiral thickening. The parenchyma
apotracheal. The secondary phloem stratified into hard (fibrous) and soft
(parenchymatous) zones. Included phloem absent. The wood commonly
partially storied (VP). Tyloses absent (?).
Reproductive type, pollination. Plants

hermaphrodite, or monoecious (rarely), or

dioecious.
Inflorescence, floral, fruit and seed morphology. Flowers

solitary, or aggregated in
inflorescences. The ultimate inflorescence units when flowers aggregated,
racemose. Flowers regular; cyclic, or partially acyclic. Sometimes the
androecium acyclic (spiralled). Free hypanthium absent. Hypogynous disk
present.
Perianth with distinct calyx and corolla (usually P3+3+3, with the outer one or
two whorls sepaloid); usually 9; usually 3 whorled; isomerous. Calyx 3, or

6; usually 2 whorled; polysepalous; valvate, or open in bud. Corolla 3; 12

whorled; polypetalous; imbricate, or valvate.
Androecium usually 25100 (i.e. many). Androecial members when many,
maturing centripetally; free of the perianth; all equal; free of one another; rarely 3
whorled, or 6 whorled (otherwise spiralled). Androecium exclusively of fertile
stamens (usually), or including staminodes (e.g. in Uvaria spp., where the outer
members may be imperfect). Staminodes when present, external to the fertile
stamens; non-petaloid. Stamens 25100 (many). Anthers adnate; nonversatile; dehiscing via longitudinal slits, or dehiscing by longitudinal valves;
extrorse; tetrasporangiate; appendaged (via expansion of the connective).
Endothecium developing fibrous thickenings. Microsporogenesis successive, or
simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or
decussate. Anther wall initially with more than one middle layer; of the dicot
type. Tapetum glandular. Pollen monosiphonous; shed in aggregates (5 genera),
or shed as single grains; when aggregated, in tetrads (usually), or in polyads
(octads in Trigynaea). Pollen grains aperturate (usually), or nonaperturate; 1
aperturate (mostly), or 2 aperturate; (mono) sulcate (mostly, occasionally with
two parallel furrows at the equator), or ulcerate (rarely); 2-celled.
Gynoecium usually 10100 carpelled (or more i.e. many). The pistil when
syncarpous, 1 celled, or 215 celled (or more?). Gynoecium apocarpous
(usually), or syncarpous (rarely); eu-apocarpous (the carpels spiralled or cyclic),
or synstylovarious to eu-syncarpous (e.g. Monodora); superior. Carpel 110
ovuled. Placentation of free carpels basal. Ovary when syncarpous 1 locular, or
215 locular (or more i.e. multilocular). Stigmas wet type; papillate; Group
III type. Placentation when unilocular parietal, or basal; when plurilocular basal.
Ovules in the single cavity when unilocular, 150; 1050 per locule (i.e. many);
ascending; apotropous; with ventral raphe; arillate, or non-arillate; anatropous;
bitegmic; crassinucellate. Outer integument not contributing to the micropyle.
Endothelium not differentiated. Embryo-sac developmentPolygonum-type. Polar
nuclei fusing prior to fertilization, or fusing only after one has been fertilized (?).
Antipodal cells formed; 3; not proliferating; ephemeral. Synergids pear-shaped.
Hypostase present, or absent. Endosperm formation cellular. Embryogeny
onagrad.
Fruit fleshy; an aggregate (commonly an aggregate of berries). The fruiting
carpels coalescing into a secondary syncarp, or not coalescing. The fruiting
carpel indehiscent; baccate. Seeds endospermic. Endospermruminate (by
contrast with Magnoliaceae); oily. Seeds with amyloid. Embryo
achlorophyllous (4/5).
Seedling. Germination

phanerocotylar, or cryptocotylar.

Physiology, phytochemistry. Anatomy

non-C type (Annona). Sugars transported

as sucrose (usually), or as oligosaccharides + sucrose (Annona squamosa).
Cyanogenic, or not cyanogenic. Alkaloids present (commonly), or absent.
Arbutin absent. Iridoids not detected. Proanthocyanidins present, or absent; when
present, cyanidin. Flavonols present, or absent; when present, quercetin. Ellagic
acid absent (2 genera, 3 species). Sieve-tube plastids P-type; type I (a).
Geography, cytology. Sub-tropical

and tropical. Widespread, especially Old

World. X = 7, 8, 9.
Taxonomy. Subclass

Dicotyledonae; Crassinucelli. Dahlgrens Superorder

Magnoliiflorae; Annonales. Cronquists Subclass Magnoliidae; Magnoliales.
APG 3 core angiosperms; Superorder Magnolianae; Order Magnoliales.

Species 1200. Genera

126; Afroguatteria, Alphonsea, Ambavia, Anaxagorea, Ancana, Annickia, Annon
a, Anomianthus, Anonidium, Artabotrys, Asimina, Asteranthe, Balonga, Bocagea,
Bocageopsis, Boutiquea,Cananga, Cardiopetalum, Chieniodendron, Cleistochla
mys, Cleistopholis, Cremastosperma, Cyathocalyx, Cyathostemma, Cymbopetalu
m, Dasoclema, Dasymaschalon, Deeringothamnus, Dendrokingstonia, Dennettia,
Desmopsis, Desmos, Diclinanona, Dielsiothamnus, Disepalum, Duckeanthus, Du
guetia, Ellipeia, Ellipeiopsis, Enicosanthum, Ephedranthus, Exellia, Fissistigma,
Fitzalania, Friesodielsia, Froesiodendron, Fusaea,Gilbertiella, Goniothalamus,
Greenwayodendron, Guamia, Guatteria, Guatteriella, Guatteriopsis, Haplostich
anthus, Heteropetalum, Hexalobus, Hornschuchia, Isolona, Letestudoxa, Lettowi
anthus, Malmea,Marsypopetalum, Meiocarpidium, Meiogyne, Melodorum, Mezz
ettia, Mezzettiopsis, Miliusa, Mischogyne, Mitrella, Mitrephora, Mkilua, Monant
hotaxis, Monocarpia, Monocyclanthus, Monodora, Neostenanthera, Neo-uvaria,
Oncodostigma, Onychopetalum, Ophrypetalum, Oreomitra,Orophea, Oxandra, P
achypodanthium, Papualthia, Petalolophus, Phaeanthus, Phoenicanthus, Piptost
igma, Platymitra, Polyalthia, Polyaulax,Polyceratocarpus, Popowia, Porcelia, P
seudartabotrys, Pseudephedranthus, Pseudoxandra, Pseuduvaria, Pyramidanthe,
Raimondia, Reedrollinsia, Richella, Rollinia, Ruizodendron, Sageraea, Sapranth
us,Schefferomitra, Sphaerocoryne, Stelechocarpus, Stenanona, Tetrameranthus,
Tetrapetalum, Toussaintia, Tridimeris, Trigynaea, Trivalvaria, Unonopsis, Uvari
a, Uvariastrum, Uvariodendron, Uvariopsis,Woodiellantha, Xylopia.
Economic uses, etc. Important

commercial fruits from Annona spp. (atemoya,

cherimoya, custard-apple, ilarma, sugar-apple, sweet sop, sour sop), Artabotrys.