The Moraceae — often called the mulberry family or fig family — are a family of

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

s comprising about 38 genera and over 1100 species. Most are widespread in tropical and subtropical regions, less so in temperate climates; however, their distribution is cosmopolitan overall. The only

synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...

within the Moraceae is presence of laticifers and milky

sap Sap is a fluid transported in xylem cells (vessel elements or tracheids) or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant. Sap is distinct from latex, resin, or cell sap; it is a separ ...

in all parenchymatous tissues, but generally useful field characters include two

carpels Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...

sometimes with one reduced, compound inconspicuous

flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...

s, and compound

fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...

s. The family includes well-known plants such as the

fig The fig is the edible fruit of ''Ficus carica'', a species of small tree in the flowering plant family Moraceae. Native to the Mediterranean and western Asia, it has been cultivated since ancient times and is now widely grown throughout the world ...

banyan A banyan, also spelled "banian", is a fig that develops accessory trunks from adventitious prop roots, allowing the tree to spread outwards indefinitely. This distinguishes banyans from other trees with a strangler habit that begin life as a ...

breadfruit Breadfruit (''Artocarpus altilis'') is a species of flowering tree in the mulberry and jackfruit family (Moraceae) believed to be a domesticated descendant of ''Artocarpus camansi'' originating in New Guinea, the Maluku Islands, and the Philippi ...

jackfruit The jackfruit (''Artocarpus heterophyllus''), also known as jack tree, is a species of tree in the fig, mulberry, and breadfruit family (Moraceae). Its origin is in the region between the Western Ghats of southern India, all of Bangladesh, ...

mulberry ''Morus'', a genus of flowering plants in the family Moraceae, consists of diverse species of deciduous trees commonly known as mulberries, growing wild and under cultivation in many temperate world regions. Generally, the genus has 64 identif ...

, and

Osage orange ''Maclura pomifera'', commonly known as the Osage orange ( ), is a small deciduous tree or large shrub, native to the south-central United States. It typically grows about tall. The distinctive fruit, a multiple fruit, is roughly spherical ...

. The 'flowers' of Moraceae are often pseudanthia (reduced

inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...

s).

Historical taxonomy

Formerly included within the now defunct order

Urticales Urticales is an order of flowering plants. Before molecular phylogenetics became an important part of plant taxonomy, Urticales was recognized in many, perhaps even most, systems of plant classification, with some variations in circumscription. A ...

, recent molecular studies have resulted in the family's placement within the

Rosales Rosales () is an order of flowering plants. Peter F. Stevens (2001 onwards). "Rosales". At: Trees At: Angiosperm Phylogeny Website. At: Missouri Botanical Garden Website. (see ''External links'' below) It is sister to a clade consisting of Faga ...

in a clade called the urticalean rosids that also includes

Ulmaceae The Ulmaceae () are a family of flowering plants that includes the elms (genus ''Ulmus''), and the zelkovas (genus ''Zelkova''). Members of the family are widely distributed throughout the north temperate zone, and have a scattered distribution ...

Celtidaceae ''Celtis'' is a genus of about 60–70 species of deciduous trees, commonly known as hackberries or nettle trees, widespread in warm temperate regions of the Northern Hemisphere. The genus is part of the extended hemp family (Cannabaceae). De ...

Cannabaceae Cannabaceae is a small family of flowering plants, known as the hemp family. As now circumscribed, the family includes about 170 species grouped in about 11 genera, including ''Cannabis'' (hemp), '' Humulus'' (hops) and '' Celtis'' (hackberries ...

, and

Urticaceae The Urticaceae are a family, the nettle family, of flowering plants. The family name comes from the genus ''Urtica''. The Urticaceae include a number of well-known and useful plants, including nettles in the genus ''Urtica'', ramie (''Boehmeri ...

. ''

Cecropia ''Cecropia'' is a Neotropical genus consisting of 61 recognized species with a highly distinctive lineage of dioecious trees. The genus consists of pioneer trees in the more or less humid parts of the Neotropics, with the majority of the speci ...

'', which has variously been placed in the Moraceae, Urticaceae, or their own family, Cecropiaceae, is now included in the Urticaceae.

Dioecy Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...

(having individuals with separate sexes) appears to be the primitive state in Moraceae.

Monoecy Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system alongside gynomonoecy, andromonoecy and trimonoecy. Monoecy is conne ...

has evolved independently at least four times within the family.

Characteristics

Flowers

The individual flowers are often small, with single whorled or absent

perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower, and structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepals when ...

. Most flowers have either petals or sepals, but not both, known as monochlamydeae, and have pistils and stamens in different flowers, known as

diclinous Plant reproductive morphology is the study of the physical form and structure (the morphology) of those parts of plants directly or indirectly concerned with sexual reproduction. Among all living organisms, flowers, which are the reproductive s ...

. Except for '' Brosimum gaudichaudii'' and ''

Castilla elastica ''Castilla elastica'', the Panama rubber tree, is a tree native to the tropical areas of Mexico, Central America, and northern South America. It was the principal source of latex among the Mesoamerican peoples in pre-Columbian times. The latex g ...

'', the perianth in all species of the Moraceae contain sepals. If the flower has an inflexed stamen, then pollen is released and distributed by wind dispersal; however, if the stamen is straight, then insect pollination is most likely to occur. Insect pollination occurs in '' Antiaropsis'', ''

Artocarpus ''Artocarpus'' is a genus of approximately 60 trees and shrubs of Southeast Asian and Pacific origin, belonging to the mulberry family, Moraceae. Most species of ''Artocarpus'' are restricted to Southeast Asia; a few cultivated species are more w ...

'', '' Castilla'', ''

Dorstenia ''Dorstenia'' is a genus within the mulberry family, Moraceae. Depending on the author, there are said to be 100 to 170 species within this genus, second only in number to the genus ''Ficus'' within Moraceae. ''Dorstenia'' species are mainly kno ...

'', ''

Ficus ''Ficus'' ( or ) is a genus of about 850 species of woody trees, shrubs, vines, epiphytes and hemiepiphytes in the family Moraceae. Collectively known as fig trees or figs, they are native throughout the tropics with a few species extending in ...

'', and '' Mesogyne''.

Leaves

The leaves are much like the flowers when analyzing diversity. The leaves can be singly attached to the stem or alternating, they may be lobed or unlobed, and can be evergreen or deciduous depending on the species in question. The

red mulberry ''Morus rubra'', commonly known as the red mulberry, is a species of mulberry native to eastern and central North America. It is found from Ontario, Minnesota, and Vermont south to southern Florida, and west as far as southeastern South Dakota, ...

can host numerous leaf types on the same tree. Leaves can be both lobed and unlobed and appear very different, but coexist on the same plant.

Fruits and seeds

Plant species in the Moraceae are best known for their fruits. Overall, most species produced a fleshy fruit containing seeds. Examples include the

from ''Artocarpus altillis'', the

from ''Morus rubra'', the

from ''Ficus carica'', and the

from ''Artocarpus heterophyllus''.

Distribution

Moraceae can be found throughout the world with a cosmopolitan distribution, thought to be due to the breakup of

Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...

during the

Jurassic period The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...

. The majority of species originate in the Old World tropics, particularly in Asia and the Pacific islands.

Phylogeny

Modern

molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

suggest these relationships:

Tribes and genera

The Moraceae comprise: *

Artocarpeae Artocarpeae is a tribe (biology), tribe within the plant family Moraceae. It includes 7 to 12 genus, genera and 70 to 87 species including ''Artocarpus altilis'', the breadfruit. Description Species in the Artocarpeae are tropical trees or shru ...

Lam. & DC. 1806 ** ''

'' J.R.Forst. & G.Forst. (c. 50 spp.) ** '' Batocarpus'' H.Karst. (4 spp.) ** '' Clarisia'' Ruiz & Pav. (3 spp.) ** '' Hullettia'' King ex Hook.f. (2 spp.) ** '' Parartocarpus'' Baill. (3 spp.) ** ''

Prainea ''Prainea'' is a genus of trees in the plant family Moraceae that is native to Southeast Asia. It is sometimes treated as a subgenus of Artocarpus. It is dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that i ...

'' King ex Hook.f. (4 spp.) ** ''

Treculia ''Treculia'' is a genus of trees in the plant family Moraceae that is native to west and central Africa and Madagascar. The best-known member of the genus, ''Treculia africana'', commonly known as the African breadfruit, is used as a food plant. ...

'' Decne. ex Trécul (3 spp.) *

Castilleae The Castilleae are a tribe within the plant family Moraceae. It includes eight to 11 genera and 55–60 species including '' Castilla'', the Panama rubber tree. Members of the tribe are primarily Neotropical with two Afrotropical genera, one g ...

C.C.Berg 1977 ** Antiaropsineae (C.C.Berg) Clement & Weiblen 2005 *** '' Antiaropsis'' K.Schum. (1 sp.) *** '' Sparattosyce'' Bur. (1 sp.) ** Castillineae Clement & Weiblen 2009 *** ''

Antiaris ''Antiaris toxicaria'' is a tree in the mulberry and fig family, Moraceae. It is the only species currently recognized in the genus ''Antiaris''. The genus ''Antiaris'' was at one time considered to consist of several species, but is now regarde ...

'' Lesch. (1 sp.) *** '' Castilla'' Cerv. (3 spp.) *** '' Helicostylis'' Trécul (7 spp.) *** ''

Maquira ''Maquira'' is a genus of trees in the family Moraceae, native to South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern He ...

'' Aubl. (5 spp.) *** '' Mesogyne'' Engl. (1 sp.) *** ''

Naucleopsis ''Naucleopsis'' is a plant genus in family Moraceae The Moraceae — often called the mulberry family or fig family — are a family of flowering plants comprising about 38 genera and over 1100 species. Most are widespread in tropical and subt ...

'' Miq. (c. 20 spp.) *** ''

Perebea ''Perebea'' is a genus of plant in family Moraceae. Selected species * '' P. angustifolia'' (Poepp. & Endl.) C.C.Berg * '' P. glabrifolia'' (Ducke) C.C.Berg. * '' P. guianensis'' Aubl. ( type) * '' P. hispidula'' Standl. * '' ...

'' Aubl. (9 spp.) *** '' Poulsenia'' Eggers (1 sp.) *** ''

Pseudolmedia ''Pseudolmedia'' is a flowering plant genus in the mulberry family (biology), family (Moraceae). Species are found in southern Mexico, the Caribbean, and Mesoamerica, Meso- and South America. They are known in Latin America as ''lechechiva'' (app ...

'' Trécul (c. 9 spp.) * Dorstenieae Dumort. 1830 ** '' Bleekrodea'' Blume (3 spp.) ** '' Bosqueiopsis'' De Wild. & T.Durand (1 sp.) ** ''

Brosimum ''Brosimum'' is a genus of plants in the family Moraceae, native to tropical regions of the Americas. The breadnut ('' B. alicastrum'') was used by the Maya civilization for its edible nut. The dense vividly colored scarlet wood of '' B. par ...

'' Sw. (13 spp.) ** ''

Broussonetia ''Broussonetia'' is a genus of four species (including one hybrid species) of trees in the family Moraceae, native to eastern Asia. These four species have high-quality fiber which consist of more than 90% of cellulose. They are traditionally ap ...

'' L’Hér. ex Vent. (8 spp.) ** ''

'' L. (c. 105 spp.) ** ''

Fatoua ''Fatoua'' is a genus of 3 species of herbs in the mulberry family. Species *'' Fatoua villosa'' *'' Fatoua pilosa'' *'' Fatoua madagascariensis'' Bibliography {{Taxonbar, from=Q147612 Moraceae genera Moraceae ...

'' Gaudich. (3 spp.) ** '' Helianthostylis'' Baill. (2 spp.) ** '' Malaisia'' Blanco 1837 (1 sp.) ** '' Scyphosyce'' Baill. (2 spp.) ** '' Sloetia'' Teijsm. & Binn. ex Kurz 1864 (1 sp.) ** '' Trilepisium'' Thouars (1 sp.) ** '' Trymatococcus'' Poepp. & Endl. (2 spp.) ** '' Utsetela'' Pellegr. (1 sp.) * Ficeae Gaudich. 1830 **''

'' L. (750 spp.) * Maclureae W.L. Clement & Weiblen 2009 ** ''

Maclura ''Maclura'' is a genus of flowering plants in the mulberry family, Moraceae. It includes the inedible Osage orange, which is used as mosquito repellent and grown throughout the United States as a hedging plant. It is dioecious, with male and fe ...

'' Nutt. (11 spp.) * Moreae Dumort. 1829 ** ''

Bagassa ''Bagassa guianensis'' is a tree in the plant family Moraceae which is native to the Guianas and Brazil. It is valued as a timber tree and as a food tree for wildlife. The juvenile leaves are distinctly different in appearance from the mature l ...

'' Aubl. (1 sp.) ** '' Milicia'' Sim (2 spp.) ** '' Morus'' L. (c. 12 spp.) ** ''

Sorocea ''Sorocea'' is a Neotropical genus of woody plants in the family Moraceae. Its distribution ranges from Chiapas to southern Brazil. It is placed within the tribe Moreae, and is closely related to the Monotypic taxon, monotypic ''Bagassa''. Spec ...

'' A.St.-Hil. (22 spp.) ** '' Streblus'' Lour. (c. 24 spp.) ** ''

Trophis ''Trophis'' is a genus in the plant family Moraceae which includes about nine species, six of which are Neotropical and three which are Palaeotropical. It is dioecious, with male and female flowers borne on separate plants. Recent work suggest ...

'' P.Browne (8 spp.) *Fossils ** †'' Artocarpoides''

References

External links

Moraceae
i
L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants.
{{Authority control Rosales Rosid families Extant Campanian first appearances