Moraceae Link

Including Artocarp(ac)eae R.Br.

Habit and leaf form. Trees, or shrubs, or lianas (including `stranglers'), or herbs (a few); laticiferous (usually), or with coloured juice (Malaisia, Fatoua); resinous, or not resinous. Perennial. Self supporting, or epiphytic, or climbing. Mesophytic. Heterophyllous, or not heterophyllous. Leaves evergreen; medium-sized to large; alternate, or opposite; when alternate, spiral, or distichous; leathery, or `herbaceous'; petiolate; non-sheathing; simple; epulvinate. Lamina dissected, or entire; when dissected, pinnatifid, or palmately lobed; pinnately veined, or palmately veined; cross-venulate. Leaves variously stipulate. Stipules interpetiolar, or intrapetiolar; free of one another, or concrescent; ochreate, or not ochreate; sometimes minute; caducous (often), or persistent. Leaves without a persistent basal meristem. Domatia recorded (2 genera); represented by pockets, or hair tufts.

General anatomy. Plants with laticifers (usually, non-articulated and branched), or without laticifers (rarely).

Leaf anatomy. Hydathodes present (occasionally), or absent. Mucilaginous epidermis present, or absent.

Adaxial hypodermis present, or absent. Cystoliths very commonly present. The mesophyll with sclerencymatous idioblasts, or without sclerenchymatous idioblasts. Minor leaf veins without phloem transfer cells (Ficus, Maclura).

Stem anatomy. Secretory cavities present, or absent (?); with latex, or with mucilage. Cork cambium present; initially superficial. Nodes tri-lacunar, or penta-lacunar. 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, or not stratified. `Included' phloem absent. Xylem with fibre tracheids, or without fibre tracheids; with libriform fibres, or without libriform fibres; with vessels. Vessel end-walls simple. Vessels without vestured pits. Wood storied, or partially storied; parenchyma typically paratracheal. Sieve-tube plastids S-type (with or without starch).

Reproductive type, pollination. Monoecious, or dioecious. Anemophilous, or entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers tightly aggregated in `inflorescences', or solitary (rarely); in spikes, in heads, and in umbels (etc.). The terminal inflorescence unit cymose. Inflorescences axillary; mostly with the small flowers packed into spikes, in heads or hollowed receptacles, on disks, etc.; with involucral bracts, or without involucral bracts; pseudanthial, or not pseudanthial. Flowers small (reduced); regular; cyclic.

Perianth sepaline (usually), or vestigial to absent (rarely); 0 (rarely), or (1-)4-5, or 8 (rarely); more or less joined; 1 whorled, or 2 whorled. Calyx (1-)4-5(-8); gamosepalous (basally connate); entire, or lobulate, or lobed; regular; persistent; imbricate, or valvate.

Androecium 1-4(-8) (in male flowers). Androecial members free of the perianth; free of one another. Androecium exclusively of fertile stamens. Stamens 1-4(-8); isomerous with the perianth (usually), or reduced in number relative to the adjacent perianth (10%); oppositisepalous (usually isostemonous); erect in bud (Moreae), or inflexed in bud. Anthers dehiscing via longitudinal slits (not dehiscing explosively); extrorse, or introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with more than one middle layer (2 to 5); of the `monocot' type. Tapetum glandular. Pollen grains aperturate; 2-4(-5) - aperturate; porate; 2-celled.

Gynoecium in female flowers 2(-3) (1-2 usually abortive). Carpels reduced in number relative to the perianth. Gynoecium syncarpous; synovarious to synstylovarious; superior to partly inferior. Ovary 1(-2) locular. Gynoecium stylate. Styles 2; free to partially joined; apical, or lateral. Stigmas 2; dry type; papillate, or non-papillate; Group II type. Placentation apical (usually, the ovule pendulous), or basal (rarely, the ovule then erect). Ovules in the single cavity 1; pendulous (usually), or ascending (rarely, when basal); anatropous to campylotropous, or hemianatropous; bitegmic; crassinucellate. Outer integument not contributing to the micropyle. Embryo-sac development Polygonum-type, or Allium-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; proliferating (Dorstenia), or not proliferating; ephemeral, or persistent (Dorstenia). Synergids hooked. Hypostase present, or absent. Endosperm formation nuclear. Embryogeny asterad (or remaining at the octant stage in Ficus).

Fruit fleshy, or non-fleshy; indehiscent; a drupe, or achene-like; enclosed in the fleshy receptacle (often, and the inflorescence axis often constituting a common fleshy receptacle), or without fleshy investment (?). Fruits from adjoining flowers aggregated into compound fruits (frequently), or not aggregated into compound fruits. The fruit aggregates coalescing (often, these sometimes explosively discharging the `seeds'), or not coalescing. Dispersal unit the seed, or the fruit, or the inflorescence. Fruit 1 seeded. Seeds endospermic, or non-endospermic. Endosperm oily, or not oily. Cotyledons 1 (by suppression), or 2. Embryo chlorophyllous (Streblus asper), or achlorophyllous (7/8); straight, or curved.

Seedling. Germination phanerocotylar, or cryptocotylar.

Physiology, biochemistry. Cyanogenic, or not cyanogenic (mostly). Alkaloids present (rarely), or absent. Iridoids absent. Proanthocyanidins present, or absent; when present, cyanidin (Ficus). Flavonols present, or absent; quercetin, or kaempferol and quercetin. Ellagic acid absent (13 species, 6 genera). Arbutin absent. Saponins/sapogenins present, or absent. Aluminium accumulation not found. C3. C3 recorded in Ficus, Morus. Anatomy non-C4 type (Artocarpus, Ficus).

Geography, cytology. Temperate (a few), or temperate to sub-tropical (most). Widespread in tropical, subtropical and warm regions. X = 7-many.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren's Superorder Malviflorae; Urticales. Cronquist's Subclass Hamamelidae; Urticales. Takhtajan's Subclass Hamamelidae; Hamamelidanae; Urticales. Species 1400. Genera about 40; Antiaris, Antiaropsis, Artocarpus, Bagassa, Batocarpus, Bosqueiopsis, Brosimum, Broussonetia, Castilla, Clarisia, Craterogyne, Dorstenia, Fatoua, Ficus, Helianthostylis, Helicostylis, Hullettia, Maclura, Maquira, Mesogyne, Metatrophis, Milicia, Morus, Naucleopsis, Olmedia (= Trophis), Olmediopsis, Parartocarpus, Perebia, Poulsenia, Prainea, Pseudolmedia, Scyphosyce, Sorocea, Sparattosyce, Streblus, Treculia, Trilepisium, Trophis, Trymatococcus, Utsetela.

Economic uses, etc. Economically important for many edible `fruits', e.g. figs (Ficus), mulberry (Morus), breadfruit and jackfruit (Artocarpus).

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