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The families of flowering plants

L. Watson and M.J. Dallwitz

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, somewhat woody); 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 persistent; 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. Leaf development not ‘graminaceous’. Domatia occurring in the family (known from 2 genera); manifested as pockets, or hair tufts.

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

Leaf anatomy. The leaf lamina dorsiventral (usually), or bifacial (the mesophyll occasionally being entirely palisade). Leaves with ‘pearl glands’ (sometimes), or without ‘pearl glands’. Hydathodes present (occasionally), or absent. Mucilaginous epidermis present, or absent. Hairs present; of numerous forms, eglandular and glandular. Adaxial hypodermis present, or absent. Lamina with secretory cavities, or with secretory cavities. Secretory cavities containing mucilage. Cystoliths very commonly present (especially in epidermes, variable in size and shape), or absent (with silicified and calcified cell walls also common). The mesophyll with sclerenchymatous idioblasts, or without sclerenchymatous idioblasts; containing crystals. The crystals druses and solitary-prismatic. Minor leaf veins without phloem transfer cells (Ficus, Maclura).

Axial (stem, wood) anatomy. Secretory cavities present, or absent; with mucilage, or with latex. Cork cambium present; initially superficial. Nodes tri-lacunar, or penta-lacunar. Primary vascular tissues in a cylinder, without separate bundles, or comprising a ring of bundles (usually traversed by wide primary rays, which sometimes broaden in the phloem as the secondary cylinder develops); collateral. Internal phloem absent. Cortical bundles absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring.

The wood ring porous to diffuse porous. The vessels large (usually medium sized); variously arranged, but often prediminantly solitary. The vessel end-walls simple. The vessels without vestured pits; with spiral thickening, or without spiral thickening. The axial xylem with fibre tracheids to without fibre tracheids; with libriform fibres (the fibres typically with small simple pits, but sometimes the pits exhibit slight borders); at least sometimes including septate fibres, or without septate fibres. The fibres without spiral thickening. The parenchyma typically paratracheal. The secondary phloem stratified into hard (fibrous) and soft (parenchymatous) zones, or not stratified. ‘Included’ phloem absent. The wood storied to partially storied, or not storied. Tyloses present, or absent.

Reproductive type, pollination. Fertile flowers functionally male and functionally female, or functionally male, or functionally female. Plants monoecious, or dioecious. Pollination anemophilous, or entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers tightly aggregated in ‘inflorescences’, or solitary (rarely); when aggregated, in spikes, in heads, and in umbels (etc.). The ultimate inflorescence units 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 when present, (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 (in 4 genra).

Gynoecium in female flowers 2(–3) carpelled (one or two of them 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 external to the original ovary (?). 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.

Seedling. Germination phanerocotylar, or cryptocotylar.

Physiology, phytochemistry. C3. C3 physiology recorded directly in Ficus, Morus. Anatomy non-C4 type (Artocarpus, Ficus). Sugars transported as sucrose, or as sugar alcohols + oligosaccharides + sucrose. Cyanogenic, or not cyanogenic (mostly). Alkaloids present (rarely), or absent. Arbutin absent. Iridoids not detected. Saponins/sapogenins present, or 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). Aluminium accumulation not found. Sieve-tube plastids S-type (with or without starch).

Geography, cytology. Temperate (a few), 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 III core angiosperms; core eudicot; Superorder Rosanae; fabid. APG IV Order Rosales.

Species 1400. Genera about 40; Antiaris, Antiaropsis, Artocarpus, Bagassa, Batocarpus, Bosqueiopsis, Brosimum, Broussonetia, Castilla, Clarisia, Craterogyne, Cudrania (= Maclura), 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).


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)

Illustrations. • Le Maout and Decaisne: Antiaris, Dorstenia, Ficus, Morus, Maclura, Sorocea. • Artocarpus altilis (as A. incisa): Lindley. • Morus alba: Lindley. • Castilla tunu, as Castilloa: Hook. Ic. Pl. 27 (1900). • Dorstenia cayapia sub-sp. asaroides, as D. asaroides: Hook. Ic. Pl. 4 (1841). • Ficus sarmentosa var. henryi, as F. foveolata: Hook. Ic. Pl. 19 (1889). • Maclura africana, as Cardiogyne: Hook. Ic. Pl. 25 (1896). • Scyphosyce manniana: Hook. Ic. Pl. 14 (1880–82). • TS leaves of Ficus elastica, F. carica and Broussonetia papyrifera, showing cystoliths: Solereder, 1908).

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 5th March 2018.’.