Unique Type of Angiosperm Pollen from the Family

Annonaceae
James W. Walker 1
1
Depatrment of Botany, University of Masschusetts, Amherst 01002

The primitive antgiosperm family Annonaceae possesses a remarkable type of pollen that appears
unique in its size (diameter up to 350 micrometers), lack of exine over nearly 50 percent of the
grain surface at maturity, and proximalipolar aperture. This unique pollent is further
distinguished by being in polyads which are compartmentalized individually within septate
stamens.

ANNONACEAE
HABIT: Characterized by the total absence of herbs. Plants may be trees (Poly.), sometimes shrubs (Ann.); rarely climbers.

LEAVES: Simple, entire, alternate, distichous, exstipulate; often aromatic and gland-dotted.

INFLORESCENCE: Axillary, sometimes on old wood (cauliflorous);solitary or in umbels or fasicle.

FLOWERS: Fairly large, generally greenish, actinomorphic, bisexual, hypogynous, trimerous.

F.F. :

CALYX: Polysepalous, of 3 sepals which may be united at the base and valvate.

COROLLA: Polypetalous, of 6 petals in 2 whorls of 3 each, of which the inner may be greatly reduced or completely absent as in Annona; Valvate or
imbricate, thick and fleshy.

ANDROECIUM: of an indefinite number of free stamens which are spirally arranged on a convex thalamus; filaments short or absent; anthers extrose;
connective produced beyond the antherlobes.

GYNOECIUM: of an indefinite number of free carpels spirally arranged on the convex thalamus; ovary superior; each ovary 1 chambered with ovules in a
double row on marginal placenta, sometimes on basal placenta; style short or absent.

FRUIT: Etaerio of berries or a fleshy berry (Ann.)

SEEDS: Large, often arillate with plenty of endosperm which is deeply ruminated.

1. Annona squamosa - Custard apple

2. Annona reticulata - bullocks heart
3. Annona muricata -
4. Artabotrys odoratissimus - Green champa
5. Cananga odorata - oil extracted from flowers used in medicine and perfumery.
6. Polyalthia longifolia - Mast tree (var. pendulata)
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; non-sheathing; 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 (in 3 genera);
manifested as pockets (usually), or hair tufts (in 1/12 species).

Leaf anatomy. Abaxial epidermis papillose, or not papillose. Stomata paracytic.

Adaxial hypodermis absent. Lamina dorsiventral; 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 sclerencymatous idioblasts, or without sclerenchymatous
idioblasts. Main veins vertically transcurrent. Minor leaf veins without phloem transfer cells
(Annona).

Stem anatomy. Secretory cavities present; with resin. Cork cambium present; initially
superficial. Nodes unilacunar, or bilacunar (according to Lammers et al 1986). Primary vascular
tissue centrifugal. Cortical bundles absent. Medullary bundles absent. Internal phloem absent.
Secondary thickening developing from a conventional cambial ring. The secondary phloem
stratified into hard (fibrous) and soft (parenchymatous) zones. ‘Included’ phloem absent. Xylem
with fibre tracheids; with vessels. Vessel end-walls horizontal; simple. Vessels without vestured
pits. Primary medullary rays wide, or mixed wide and narrow, or narrow. Wood diffuse porous;
partially storied (VP); parenchyma apotracheal. Sieve-tube plastids P-type; type I (a). Pith with
diaphragms (commonly), or without diaphragms.

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 unit 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; 1–2 whorled; polypetalous; imbricate, or
valvate.
Androecium usually 25–100 (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 25–100 (‘many’). Anthers
adnate; non-versatile; 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 10–100 carpelled (or more — i.e. ‘many’). The pistil when syncarpous, 1
celled, or 2–15 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 1–10 ovuled. Placentation of free carpels basal. Ovary when
syncarpous 1 locular, or 2–15 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, 1–50; 10–50 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
development Polygonum-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. Endosperm ruminate (by contrast with Magnoliaceae); oily.
Seeds with amyloid. Embryo achlorophyllous (4/5).

Seedling. Germination phanerocotylar, or cryptocotylar.

Physiology, biochemistry. Cyanogenic, or not cyanogenic. Alkaloids present (commonly), or

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).
Arbutin absent. Sugars transported as sucrose (usually), or as oligosaccharides + sucrose
(Annona squamosa). Anatomy non-C4 type (Annona).

Geography, cytology. Sub-tropical and tropical. Widespread, especially Old World. X = 7, 8, 9.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Magnoliiflorae;

Annonales. Cronquist’s Subclass Magnoliidae; Magnoliales. APG (1998) basal order. APG 3
(2009) Order: Magnoliales.
Species 1200. Genera 126; Afroguatteria, Alphonsea, Ambavia, Anaxagorea, Ancana, Annickia,
Annona, Anomianthus, Anonidium, Artabotrys, Asimina, Asteranthe, Balonga, Bocagea,
Bocageopsis, Boutiquea, Cananga, Cardiopetalum, Chieniodendron, Cleistochlamys,
Cleistopholis, Cremastosperma, Cyathocalyx, Cyathostemma, Cymbopetalum, Dasoclema,
Dasymaschalon, Deeringothamnus, Dendrokingstonia, Dennettia, Desmopsis, Desmos,
Diclinanona, Dielsiothamnus, Disepalum, Duckeanthus, Duguetia, Ellipeia, Ellipeiopsis,
Enicosanthum, Ephedranthus, Exellia, Fissistigma, Fitzalania, Friesodielsia, Froesiodendron,
Fusaea, Gilbertiella, Goniothalamus, Greenwayodendron, Guamia, Guatteria, Guatteriella,
Guatteriopsis, Haplostichanthus, Heteropetalum, Hexalobus, Hornschuchia, Isolona,
Letestudoxa, Lettowianthus, Malmea, Marsypopetalum, Meiocarpidium, Meiogyne, Melodorum,
Mezzettia, Mezzettiopsis, Miliusa, Mischogyne, Mitrella, Mitrephora, Mkilua, Monanthotaxis,
Monocarpia, Monocyclanthus, Monodora, Neostenanthera, Neo-uvaria, Oncodostigma,
Onychopetalum, Ophrypetalum, Oreomitra,Orophea, Oxandra, Pachypodanthium, Papualthia,
Petalolophus, Phaeanthus, Phoenicanthus, Piptostigma, Platymitra, Polyalthia, Polyaulax,
Polyceratocarpus, Popowia, Porcelia, Pseudartabotrys, Pseudephedranthus, Pseudoxandra,
Pseuduvaria, Pyramidanthe, Raimondia, Reedrollinsia, Richella, Rollinia, Ruizodendron,
Sageraea, Sapranthus, Schefferomitra, Sphaerocoryne, Stelechocarpus, Stenanona,
Tetrameranthus, Tetrapetalum, Toussaintia, Tridimeris, Trigynaea, Trivalvaria, Unonopsis,
Uvaria, 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.