Moraceae Gaudich.

First published in Gen. Pl. [Trinius] 13. 1835 (1835)nom. cons.
This family is accepted

Descriptions

George R. Proctor (2012). Flora of the Cayman Isands (Second Edition). Royal Botanic Gardens, Kew

Morphology General Habit
Trees or shrubs, rarely herbs, often with milky sap (latex); leaves alternate, entire, toothed or lobed either pinnately or palmately; stipules present
Morphology Reproductive morphology Flowers
Flowers unisexual, in spikes or heads, or on a flat entire or lobed receptacle, or on the inside of a closed receptacle; perianth sepaloid, of 2–5 (usually 4) free or fused segments Male flowers with stamens generally as many as the perianth lobes and opposite them, or only 1. Female flowers with 1-celled ovary, superior to inferior, bearing a simple, 2-toothed or 2-partite style; ovule 1
Morphology Reproductive morphology Fruits
Fruit 1-seeded, free or more commonly united with fruits of other flowers to form a more or less fleshy pseudocarp, in whose formation the receptacle may also be involved.
Distribution
A mostly tropical family of about 53 genera and more than 1,000 species.
[Cayman]

Timothy M. A. Utteridge and Laura V. S. Jennings (2022). Trees of New Guinea. Kew Publishing. Royal Botanic Gardens, Kew

Distribution
A family of 39 genera and about 1100 species from tropical and warm temperate regions worldwide. Eleven genera occur in New Guinea.
Recognition
Moraceae can be recognised by the presence of milky sap and stipules which often leave circular scars, leaves often with triplinerved or brochidodromous venation, tiny unisexual flowers in congested inflorescences, the unilocular ovary and apical placentation.
Morphology General Habit
Trees, shrubs, climbers and herbs
Morphology General
Sap present, described as milky or white latex
Morphology Leaves Stipules
Stipules present, free or fused, usually amplexicaul (sheathing stem), falling leaving a circular scar, sometimes lateral and intrapetiolar
Morphology Leaves
Leaves alternate and distichous, spirally arranged, occasionally opposite, margin entire to serrate, venation often conspicuous, usually pinnate or brochidodromous
Morphology Reproductive morphology Inflorescences
Inflorescences usually unisexual and plants dioecious, congested, spicate, racemose or capitate, often partially adnate to an expanded receptacle, or receptacle completely enclosing flowers within a spherical hollow (Ficus), involucral bracts often present. Flowers unisexual, free or adnate to receptacle
Morphology Reproductive morphology Flowers
Male flowers with 2–4(–8) perianth lobes, free or fused, or perianth absent; stamens 1–4(–7), inflexed or straight in bud; pistillode present or absent Female flowers usually with 4 perianth lobes, free or fused; ovary unilocular, free or adnate to perianth, ovule 1, usually apically attached, stigma 1, sometimes with 2 branches
Morphology Reproductive morphology Fruits
Fruit often drupaceous, free or adnate to perianth, often forming a multiple fruit with expanded, fleshy perianth and receptacle
Morphology Reproductive morphology Seeds
Seed with or without endosperm.
[TONG]

Timothy Utteridge & Gemma Bramley (2020). The Kew Tropical Plant Families Identification Handbook, Second Edition. Kew Publishing Royal Botanic Gardens, Kew

Note
Woody (rarely herbs) with white sap; stipules large often leaving circular scar. Leaves simple. Flowers unisexual, often grouped into dense complex heads; perianth 1 whorl; ovary superior; 1–2 stigmas.
Recognition
Characters of similar families: Urticaceae: no sap, often herbs, cystoliths often elongate, stigma 1. Cannabaceae: no sap, flowers not in heads, stipules small, caducous. Magnoliaceae: no sap, flowers large solitary bisexual, many stamens and tepals. Euphorbiaceae: petiole geniculate, stigmas 3, fruit breaks into 3 leaving columella.
Morphology General Habit
Shrubs, trees, lianas or hemi-epiphytes; milky white sap (may turn brown when dry)
Morphology Leaves Stipules
Stipules often large and amplexicaul, forming a cap over the bud and leaving a circular scar
Morphology Leaves
Leaves alternate or spiral; simple, rarely pinnate-compound; lamina margins entire or toothed; venation pinnate, often conspicuous and/or 3-veined from the base; cystoliths present, punctiform; hairs often present
Morphology Reproductive morphology Inflorescences
Inflorescences pendulous or erect spikes, cymes, racemes, glomerules, or condensed into receptacular inflorescences by fusion of flower parts and/ or bracts and often enclosed by an involucre; perianth often embedded in and fused with receptacle
Morphology Reproductive morphology Flowers
Flowers small, unisexual, rarely solitary; male flowers with tepals 2–6, stamens 1–4; female with tepals 2–6 or absent, often fused to ovary; stigmas 1–2; ovule 1
Morphology Reproductive morphology Fruits
Fruit drupe-like; sometimes dehiscent; often grouped into complex disklike or globose structures or condensed into a compound fruit (syncarp) by fusion of parts: infructescences may be urn-shaped with apical ostiole closed by bracts (‘figs’).
Distribution
40 genera and c. 1,100 species, mostly tropical, worldwide. Large genera Artocarpus (Old World), Ficus and Dorstenia.
Ecology
Tropical evergreen lowland forest, seasonal montane forest (often as pioneer species) or understorey in upland rain forest, hemi-epiphytes in the canopy.
Description Author
C M Wilmot-Dear
[KTROP-FIH]

Moraceae, C.C. Berg (University of Bergen). Flora of Tropical East Africa. 1989

Morphology General Habit
Trees, shrubs or herbs, dioecious or monoecious; sap milky, sometimes watery (but not turning black)
Morphology Leaves
Leaves in spirals or distichous, sometimes subopposite or subverticillate, entire or sometimes pinnately or palmately incised, stipulate
Morphology Reproductive morphology Inflorescences
Inflorescences bisexual or unisexual, spicate, globose-, clavate- or discoid-capitate, urceolate or uniflorous
Morphology Reproductive morphology Flowers
Staminate flowers with 2–6 tepals or perianth lacking; stamens 1–4 Pistillate flowers with 2–6 tepals or perianth lacking; pistil 1; ovary free or adnate to the perianth; stigmas 1–2; ovule 1, attached apically
sex Male
Staminate flowers with 2–6 tepals or perianth lacking; stamens 1–4
sex Female
Pistillate flowers with 2–6 tepals or perianth lacking; pistil 1; ovary free or adnate to the perianth; stigmas 1–2; ovule 1, attached apically
Morphology Reproductive morphology Fruits
Fruit achene-like, drupaceous (dehiscent or not), or forming a drupaceous whole with the fleshy perianth or with the fleshy receptacle as well
Morphology Reproductive morphology Seeds
Seed large without endosperm or small with endosperm; embryo various
[FTEA]

Moraceae, C. C. Berg. Flora Zambesiaca 9:6. 1991

Morphology General Habit
Trees, shrubs or herbs, dioecious or monoecious; sap milky, sometimes watery (but not turning black)
Morphology Leaves
Leaves spirally or distichously arranged, sometimes subopposite or subverticillate, entire or sometimes pinnately or palmately incised, stipulate
Morphology Reproductive morphology Inflorescences
Inflorescence bisexual or unisexual, spicate, globose, clavate- or discoid-capitate, urceolate, sometimes uniflorous
Morphology Reproductive morphology Flowers
Staminate flowers with 2–6 tepals or perianth lacking; stamens 1–4 Pistillate flowers with 2–6 tepals or perianth lacking; pistil 1; ovary free or adnate to the perianth; stigmas 1 or 2, ovule 1, apically attached
sex Male
Staminate flowers with 2–6 tepals or perianth lacking; stamens 1–4
sex Female
Pistillate flowers with 2–6 tepals or perianth lacking; pistil 1; ovary free or adnate to the perianth; stigmas 1 or 2, ovule 1, apically attached
Morphology Reproductive morphology Fruits
Fruit achene-like, drupaceous (dehiscent or not), or forming a drupaceous whole with the fleshy perianth or with the fleshy receptacle as well
Morphology Reproductive morphology Seeds
Seed large and without endosperm or small with endosperm, embryo various
[FZ]

Gemma Bramley, Anna Trias-Blasi & Richard Wilford (2023). The Kew Temperate Plant Families Identification Handbook. Kew Publishing Royal Botanic Gardens, Kew

Recognition
Characters of similar families: Urticaceae: no sap; often herbs, cystoliths often elongate; stigma 1, stamens inflexed in bud. Cannabaceae: no sap, flowers not in heads, lax inflorescences. Magnoliaceae: no sap, flowers large, solitary, bisexual, many stamens and tepals. Euphorbiaceae: petiole geniculate, stigmas usually >2, fruit breaks into 3 leaving columella. Salicaceae (Flacourtia): inconspicuous stipules, many stamens, female flowers with disk.
Morphology Leaves Stipules
Stipules persistent or caducous, often large and amplexicaul leaving a circular scar
Morphology Leaves
Leaves alternate (rarely opposite or whorled), simple, entire or toothed (sometimes lobed); venation pinnate (rarely palmate), often conspicuous, 3(–5)-veined from the base; punctiform cystoliths sometimes present
Morphology Reproductive morphology Inflorescences
Inflorescences pendulous or erect spikes, cymes, racemes, glomerules, or condensed by fusion of flower parts and/or bracts and often enclosed by an involucre; perianth often embedded in and fused with receptacle
Morphology General Habit
Plants monoecious or dioecious Shrubs, trees, lianas or hemi-epiphytes (only Fatoua villosa is herbaceous in the temperate region), sometimes spiny; milky white latex (drying brown, never black)
Morphology Reproductive morphology Flowers
Flowers small, unisexual, rarely solitary; tepals (2–)4(–6), free or fused, imbricate or valvate; male flowers with stamens equal and opposite tepals, straight or inflexed in bud, pistillode often present; female flowers with tepals often fused to ovary, sometimes absent; stigmas 1–2 usually filiform; 1(–2) locular, 1 ovule per locule; inferior to superior
Morphology Reproductive morphology Fruits
Fruit often dehiscent drupes, rarely achenes; often grouped into complex disk-like or globose structures or condensed into a syncarp by fusion of parts; specialised infructescence is a syconium; hollow, urceolate receptacle with apical ostiole closed by bracts (fig)
Morphology Reproductive morphology Seeds
Seeds 1.
Distribution
Family of 45 genera and ca. 1,200 species, cosmopolitan, mostly tropical and subtropical.
Ecology
Evergreen lowland forest, seasonal montane forest (often as pioneer species) or understorey in upland rainforest, hemi-epiphytes in the canopy.
Note
Shrubs, trees, lianas or hemi-epiphytes; white sap; stipules large often leaving circular scar. Leaves simple. Inflorescences diverse. Flowers unisexual, tepals usually 4, often grouped into dense complex heads; perianth 1 whorl; ovary superior; 1–2 stigmas.
Description Author
Alison Moore & C. M. Wilmot-Dear
[KTEMP-FIH]

Moraceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:2. 1958

Morphology General Habit
Trees or shrubs or rarely herbs, often with milky juice
Morphology Leaves
Leaves alternate, rarely opposite, simple; stipules paired, often caducous and leaving a scar
Morphology Reproductive morphology Flowers
Flowers much reduced, often in heads, disks or hollow receptacles, unisexual Female flower with superior or inferior ovary, of 2 carpels, one often abortive, usually 1-celled; styles mostly 2, filiform; ovule solitary, pendulous, rarely basal and erect
Morphology Reproductive morphology Flowers Calyx
Calyx-lobes usually 4, sometimes reduced, imbricate or valvate
Morphology Reproductive morphology Flowers Androecium
Stamens usually equal in number and opposite to the sepals; filaments inflexed or erect in bud; anthers 2-celled
sex Female
Female flower with superior or inferior ovary, of 2 carpels, one often abortive, usually 1-celled; styles mostly 2, filiform; ovule solitary, pendulous, rarely basal and erect
Morphology Reproductive morphology Fruits
Fruit a small achene, nut or drupe
Morphology Reproductive morphology Seeds
Seed with or without endosperm, often with curved embryo
[FWTA]

Ribeiro, J.E.L.S. (2011). Neotropical Moraceae.

Morphology
Description

Trees, shrubs, hemi-epiphytes or herbs and very rarely lianas; plants monoecious , dioecious or androdioecious, unarmed (except  Maclura Nutt.), latex present in all plant parts, white, brown (coffee with milk), yellow or orange. Leaves simple , alternate or rarely opposite (Bagassa Aubl.) arranged in spiral in Ficus L. and Dorstenia L. (rosulate) or distichous , at least in the reproductive branches in other genera, venation pinnate , rarely trinerved (Bagassa), brochidodromous ; stipules usually conspicuous, terminal , fully amplexicaul , partially amplexicaul to lateral , their scars forming a complete ring or, if not amplexicaul , with a scar opposite the insertion of the leaves. Inflorescences unisexual or bisexual , usually in pairs in the leaf axil or more on short shoots, forming unisexual or bisexual synflorescences (Tribe Ficeae and Castilleae), structurally complex, racemose, spicate, globose , capitate  or urceolate receptacles (= syconium, fig). Flowers unisexual, free or immersed in the receptacle , monochlamydeous or achlamydeous, staminate flowers with 2-4 (-7) free or connate tepals or tepals absent, stamens (3-) 4 (-5), or reduced to 1, filaments straight or inflexed before anthesis; pistillode generally absent; pistillate flowers with (3-) 4 (-7) free or connate tepals; ovary free or adnate to tepals, stigmas 2 or 1, ovule 1 with subapical placentation. Fruits achenes, usually associated with a fleshy perianth or other structures that become fleshy , like bracts and/or receptacle .

Distribution
Distribution in the Neotropics
  • The family is distributed throughout the Neotropics, with a major center of diversity in northern South America, specifically the Amazon rainforests, where all Neotropical genera are represented.
  • The genus Dorstenia has a concentration of species mainly in northern Central America and the Greater Antilles and in the Atlantic forests of southeastern Brazil.
  • In the genus Morus L., M. celtidifolia Kunth is distributed primarily from Mexico to Honduras and M. insignis Bureau in Central America and the Andean region of South America.
Diagnostic
Key differences from similar families
  • The presence of terminalstipule or stipule scar, associated with the presence of latex in all parts of the plant, makes Moraceae difficult to confuse with other families.  Some species of Coussapoa (Urticaceae) have aqueous whitish latex in the branches and stipules terminals, besides the usual hemi-epiphytichabit, which can be confused with species of Ficus, although waxy glandular spots in the base of blade underside do not occur in Coussapoa spp.
Useful tips for generic identification
  • Images of some genera and species of the Central Brazilian Amazon can be accessed at: http://brahms.inpa.gov.br/bol/content/Groups/PFRD/resources/PFRD_1999_Moraceae.pdf
Distinguishing characters (always present)
  • The most obvious features of the family are the presence of latex in all parts of the plant, and terminal stipules.
  • In times of high water stress, some species may not exude evident latex.
  • The stipules are small and often caducous in some species of Clarisia Ruiz & Pav. and Sorocea A.St-Hil. Even in the absence of stipules, it is possible to observe the scars left by these structures.
  • The flowers are small, tapaloid or naked; unisexual; gynoecium with one or two stigmas; ovary uniovulate and placentation subapical.
Other important characters
  • Generally woody plants, from trees to hemiepiphytes and lianas.
  • Leaves alternate.
  • Inflorescences structurally complex, racemose, spicate, globose, capitate or an urceolate receptacle (= syconium, fig).
Notable genera and distinguishing features

Bagassa: leaves opposite, venationtrinerved.

Batocarpus: canopy tree, stems and the shallow roots covered with elevated red lenticels.

Brosimum: inflorescences usually bisexual and globose, covered with peltate bracts.

Clarisa racemosa Ruiz & Pav.: canopy tree, stems and the shallow roots covered with elevated red lenticels.

Clarisa ilicifolia (Spreng.) Lanji. & Rossberg: understory treelets; leaf margin and apex spinulose. This species is often confused with Soroceaguilleminiana Gaudich., differentiated by the prominentmidrib on the upper surface of the blade and by the presence of uncinate trichomes.

Dorstenia: herbaceoushabit; inflorescence a receptacle with variable shape -  orbicular, elliptic, obovate, angulate, stellate or furcate.

Ficus subg. Urostigma sect. Americana: habit hemi-epiphyte; inflorescence a syconium; with a waxy glandular spot at the base of the midrib of the lower surface of the leaf.

Ficus subg. Pharmacosycea sect. Pharmacosycea: trees; inflorescence a syconium; with 2 waxy glandular spots in the axils of the basallateralveins of the lower surface of the leaf.

Maclura: tree or liana; often armed with axillary thorns.

Morus: leaves alternatedistichous, the margin crenate to serrate.

Sorocea: lamina is slightly to pronouncedly inequilateral; leaf margin mostly dentate to denticulate, sometimes the teeth and acumen spinulose.

Trymatococcus: leaves with globose-capitate pluricellular hairs; inflorescences bisexual.

Genera of tribe Castilleae (Castilla, Helicostylis, Maquira, Naucleopsis, Perebea, Poulsenia, Pseudolmedia): trees with the architectural model of Cook and the inflorescences usually with involucres of bracts.

General Description
Number of genera

19 genera:

Tribe Artocarpeae

Batocarpus H.Karst. (3 spp.)Clarisia Ruiz & Pav. (3 spp.)

Tribe Moreae

Bagassa Aubl. (1 spp.)Morus L. (2 spp.)Sorocea A. St-Hil. (14 spp.)Trophis P.Browne (5 spp.)

Tribe Maclureae

Maclura Nutt. (2 spp.)

Tribe Dorstenieae

Brosimum Sw. (15 spp.)Dorstenia L. (46 spp.)Helianthostylis Baill. (2 spp.)Trymatococcus Poepp. & Endl. (2 spp.)

Tribe Castilleae

Castilla Cerv. (3 spp.)Helicostylis Trécul (7 spp.)Maquira Aubl. (4 spp.)Naucleopsis Miq. (22 spp.)Perebea Aubl. (9 spp.)Poulsenia Eggers (2 spp.)Pseudolmedia Trécul (15 spp.)

Tribe Ficeae

Ficus subg. Urostigma (Gasparrini) Miq. sect. Americana Miq.(ca. 100 spp.)Ficus subg. Pharmacosycea (Miq.) Miq. sect. Pharmacosycea (ca. 20 spp.)

Notes on delimitation
  • The family Moraceae is currently placed in the order Rosales (APG III) as sister to the family Urticaceae (Sytsma et al., 2002), here including the genera Cecropia Loefl., Pourouma Aubl. and Coussapoa Aubl., previously placed in the family Cecropiaceae.
  • The family Moraceae is currently divided into six tribes: Artocarpeae, Morea, Maclureae, Dorstenieae, Castilleae and Ficeae (Ribeiro, 2007; Clement & Weiblen, 2009).
Status
  • All genera are endemic and restricted to the neotropics, with the exception of Morus, Trophis, Dorstenia and Ficus, which have a pantropical distribution pattern.
  • The genus Artocarpus J.R. & G. Foster and Morusalba L. are naturalized in the Neotropics, grown for their edible fruits.
General notes
  • The inflorescences of the family are among the most complex in Angiosperms. These are often associated with pollination systems in which plants provide breeding sites as a reward for the pollinator. For the genus Ficus entomophily is well documented where the pollination is carried out by wasps of the family Agaonidae (Chalcidoidae, Hymenoptera). The two neotropical sections Americana and Pharmacosycea are pollinated by the wasp genera Pegoscapus and Tetrapus respectively.
  • The Neotropical Castilla elastica Sessé (Sakai, 2001), the New Guinea Antiaropsis decipiens K. Shum. (Zerega et al., 2004) and probably all species of Tribe Castilleae are pollinated by thrips (Thysanoptera).
  • Although the family always have been cited, even in the recent scientific literature, as an example of a wind-pollinated group, probably only 6% of all species of the family are really anemophilous (Ribeiro, 2007).
Literature
Important literature

Berg, C.C. 1972. Olmedieae and Brosimeae. Flora Neotropica 7. Hafner Publishing Company, New York.Berg, C.C. Moraceae. 1992. In: A.R.A. Görts-van Rijn (ed.). Flora of the Guianas. vol. 21. Koenigstein, Germany. 82p.Berg, C.C. 1998b. Moraceae (excl. Ficus). In: G. Harling & L. Andersson (eds.). Flora of Ecuador. Vol. 27B. Arlöv, Sweden. 128p.Berg, C.C. 2001. Moreae, Artocarpeae, and Dorstenia (Moraceae). With introductions to the family and Ficus and with additions and corrections to Flora Neotropica Monograph 7. Flora Neotropica Monograph 83. The New York Botanical Garden, New York. p. 347.Berg, C.C. & M.E.E. Hijman. 1999. The genus Dorstenia (Moraceae). Ilicifolia 2:1-211.Berg, C.C. & X. Villavicencio. 2004. Taxonomic studies on Ficus (Moraceae) in the West Indies, extra-Amazonian Brazil, and Bolivia. Ilicifolia 4:1-132.Carauta, J.P.P. 1978. Dorstenia L. (Moraceae) do Brasil e Países Limítrofes. Rodriguesia 29(44): 52-223.Clement, W.L. & Weiblen, G.D. 2009. Morphological Evolution in the Mulberry Family (Moraceae). Systematic Botany 34(3): 530-552.Miquel, F.A.W. 1853 (1965). Urticineae. In: C.F.P. von Martius (ed.). Flora Brasiliensis vol. 4 part. 1, pp. 79-170. J. Cramer, Weinheim.Ribeiro, J.E.L.S. 2007. Studies on the phylogeny, taxonomy and evolution of reproductive characters in Moraceae Gaudich. Universidade Estadual de Campinas (Ph.D. thesis). 124p. (in Portuguese).Rohwer, J.G. 1993. Moraceae. In: K. Kubitzki, J.G. Rohwer & V. Bittrich (eds.). The Families and Genera of Vascular Plants. Vol. II. Spring-Verlag, Berlin. 438-453.Sakai, S. 2001. Thrips pollination of androdioecious Castillaelastica (Moraceae) in a seasonal tropical forest. American Journal of Botany 88(9): 1527-1534.

Sytsma, K.J.; Morawetz, J.; Pires, J.C.; Nepokroeff, M.; Conti, E.; Zjhra, M.; Hall, J.C. & Chase, M.W. 2002. Urticalean Rosids: circumscription, Rosid ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences. American Journal of Botany 89(9): 1531-1546.

Zerega, N.J.C., L.A. Moundy & G.D. Weiblen. 2004. Pollination in the New Guinea endemic Antiaropsisdecipiens (Moraceae) is mediated by a new species of thrips, Thripsantiaropsidis sp. nov. (Thysanoptera: Thripidae). International Journal of Plant Sciences 165(6):1017-1026.

[NTK]

Uses

Use
The large genus Ficus (870 species) contains many economically important fruit trees, which like Morus species, are widely cultivated.
[KTEMP-FIH]

Sources

  • Flora Zambesiaca

    • Flora Zambesiaca
    • http://creativecommons.org/licenses/by-nc-sa/3.0
  • Flora of Tropical East Africa

    • Flora of Tropical East Africa
    • http://creativecommons.org/licenses/by-nc-sa/3.0
  • Flora of West Tropical Africa

    • Flora of West Tropical Africa
    • http://creativecommons.org/licenses/by-nc-sa/3.0
  • Flora of the Cayman Islands

    • Flora of the Cayman Islands
    • http://creativecommons.org/licenses/by-nc-sa/3.0
  • Herbarium Catalogue Specimens

  • Kew Names and Taxonomic Backbone

    • The International Plant Names Index and World Checklist of Vascular Plants 2024. Published on the Internet at http://www.ipni.org and https://powo.science.kew.org/
    • © Copyright 2023 International Plant Names Index and World Checklist of Vascular Plants. http://creativecommons.org/licenses/by/3.0
  • Neotropikey

    • Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics.
    • http://creativecommons.org/licenses/by/3.0
  • Trees of New Guinea

    • Trees of New Guinea
    • http://creativecommons.org/licenses/by-nc-sa/3.0
  • The Kew Temperate Plant Families Identification Handbook

    • The Kew Temperate Plant Families Identification Handbook
    • http://creativecommons.org/licenses/by-nc-sa/3.0
  • The Kew Tropical Plant Families Identification Handbook

    • The Kew Tropical Plant Families Identification Handbook
    • http://creativecommons.org/licenses/by-nc-sa/3.0