Moraceae Link
Including Artocarpeae (Artocarpaceae) R.Br., Ficaceae (Dum.) Dum.
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. ‘Normal’ plants. 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 palmatifid; 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. Fertile flowers functionally male and functionally female, or functionally male, or functionally female. Plants 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 blunt-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) carpelled (1–2 usually abortive). Carpels reduced in number relative to the perianth. The pistil 1 celled, or 2 celled. 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 (?). The drupes with one stone. Gynoecia of adjoining flowers combining to form a multiple fruit (frequently), or not forming a multiple fruit. The multiple fruits 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; flat. Embryo chlorophyllous (Streblus asper), or achlorophyllous (7/8); straight, or curved. The radicle dorsal. Micropyle not zigzag.
Seedling. Germination phanerocotylar, or cryptocotylar.
Physiology, biochemistry. Cyanogenic, or not cyanogenic (mostly). Alkaloids present (rarely), or absent. Iridoids not detected. 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. Sugars transported as sucrose, or as sugar alcohols + oligosaccharides + sucrose. C3. C3 physiology recorded directly 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. APG (1998) Eudicot; core Eudicot; Rosid; Eurosid I; Rosales. 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).
Illustrations. • Technical details (Artocarpus). • Technical details (Antiaris, Dorstenia, Ficus, Morus, Maclura, Sorocea). • Technical details (Morus).
Quotations
And Thisby, tarrying in mulberry shade,
His dagger
drew, and died
(‘Midsummer Night’s Dream’, v., i)
Feed him
with apricocks and dewberries,
With purple grapes, green figs and mulberries
(‘Midsummer Night’s Dream’, iii., 1)
Cite this publication as: ‘L. Watson and M. J. Dallwitz (1992 onwards). The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 14th December 2000. http://biodiversity.uno.edu/delta/’. Dallwitz (1980), Dallwitz, Paine and Zurcher (1993, 1995, 2000), and Watson and Dallwitz (1991) should also be cited (see References).
