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Moraceae Temporal range: 80–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Cretaceous – Recent
Panama rubber tree (Castilla elastica)
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Rosids
Order:	Rosales
Family:	Moraceae Gaudich. (1835) nom. cons.
Genera
48; see text

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 subtropical regions, less so in temperate climates; however, their distribution is cosmopolitan overall. The only synapomorphy within the Moraceae is presence of laticifers and milky sap in all parenchymatous tissues, but generally useful field characters include two carpels sometimes with one reduced, compound inconspicuous flowers, and compound fruits. The family includes well-known plants such as the fig, banyan, breadfruit, jackfruit, mulberry, and Osage orange. The 'flowers' of Moraceae are often pseudanthia (reduced inflorescences).

Description

Overall

The family varies from colossal trees like the Indian Banyan (Ficus benghalensis) which can cover five acres (two hectares) of ground, to Dorstenia barnimiana which is a small stemless, bulbous succulent 2–5 cm in diameter that produces a single peltate leaf on a 4–15 cm petiole. These two species have an approximately one billion fold difference in weight.

Flowers

The individual flowers are often small, with single whorled or absent perianth. Most flowers have either petals or sepals, but not both, known as monochlamydeae, and have pistils and stamens in different flowers, known as diclinous. Except for Brosimum gaudichaudii and Castilla elastica, 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, Castilla, Dorstenia, Ficus, 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 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 breadfruit from Artocarpus altilis, the mulberry from Morus rubra, the fig from Ficus carica, and the jackfruit from Artocarpus heterophyllus.

Taxonomy

Formerly included within the now defunct order Urticales, recent molecular studies have resulted in the family's placement within the Rosales in a clade called the urticalean rosids that also includes Ulmaceae, Celtidaceae, Cannabaceae, and Urticaceae. Cecropia, which has variously been placed in the Moraceae, Urticaceae, or their own family, Cecropiaceae, is now included in the Urticaceae.

Dioecy (having individuals with separate sexes) appears to be the primitive state in Moraceae. Monoecy has evolved independently at least four times within the family.

Phylogeny

Modern molecular phylogenetics suggest these relationships:

Tribes and genera

Moraceae is comprised 48 genera in seven tribes.

• Artocarpeae Lam. & DC. 1806
  • Artocarpus J.R.Forst. & G.Forst. (73 spp.)
  • Batocarpus H.Karst. (3 spp.)
  • Clarisia Ruiz & Pav. (4 spp.)
• Chlorophoreae (syn. Maclureae W.L. Clement & Weiblen 2009)
  • Maclura Nutt. (13 spp.)
• Parartocarpeae
  • Hullettia King ex Hook.f. (2 spp.)
  • Parartocarpus Baill. (2 spp.)
  • Pseudostreblus Bureau (1 sp.)
• Olmedieae Trécul 1847 (syn. Castilleae)
  • Antiaris Lesch. (1 sp.)
  • Antiaropsis K.Schum. (2 spp.)
  • Castilla Cerv. (3 spp.)
  • Helicostylis Trécul (8 spp.)
  • Maquira Aubl. (4 spp.)
  • Mesogyne Engl. (1 sp.)
  • Naucleopsis Miq. (25 spp.)
  • Olmedia Ruiz & Pav. (1 sp.)
  • Perebea Aubl. (10 spp.)
  • Poulsenia Eggers (1 sp.)
  • Pseudolmedia Trécul (11 spp.)
  • Sparattosyce Bur. (2 spp.)
  • Streblus Lour. (5 spp.)
• Dorstenieae Dumort. 1830
  • Bleekrodea Blume (3 spp.)
  • Bosqueiopsis De Wild. & T.Durand (1 sp.)
  • Brosimum Sw. (19 spp.)
  • Broussonetia L’Hér. ex Vent. (4 spp.)
  • Dorstenia L. (122 spp.)
  • Fatoua Gaudich. (3 spp.)
  • Malaisia Blanco 1837 (1 sp.)
  • Scyphosyce Baill. (3 spp.)
  • Sloetia Teijsm. & Binn. ex Kurz (1 sp.)
  • Sloetiopsis Engl. (1 sp.)
  • Trilepisium Thouars (2 sp.)
  • Utsetela Pellegr. (2 sp.)
• Ficeae Gaudich. 1830
  • Ficus L. (880 spp.)
• Moreae Dumort. 1829
  • Afromorus E.M.Gardner
  • Ampalis Bojer (2 spp.)
  • Bagassa Aubl. (1 sp.)
  • Maillardia Frapp. ex Duch. (2 spp.)
  • Milicia Sim (2 spp.)
  • Morus L. (17 spp.)
  • Paratrophis Blume (12 spp.)
  • Sorocea A.St.-Hil. (22 spp.)
  • Taxotrophis Blume (6 spp.)
  • Trophis P.Browne (5 spp.)

Other genera accepted by Plants of the World Online as of April 2024:

• Allaeanthus Thwaites (4 spp.)
• Calaunia Gudzins. (1 sp.)
• Hijmania M.D.M.Vianna (4 spp.)
• Prainea King ex Hook.f. (2 spp.)
• Treculia Decne. ex Trécul (5 spp.)

Fossil genera and species

In addition to the living species, a number of fossil genera have been ascribed to the family:

• †Aginoxylon Dupéron, 1977
• Aginoxylon moroides Dupéron, 1977
• †Artocarpidium Unger, 1850
• †Artocarpoides Saporta, 1865
• †Arthmiocarpus Delevoryas, 1964
• †Artocarpoxylon Prakash & Lalitha, 1978
• †Becktonia M. Chandler, 1961
  • †Becktonia hantonensis M. Chandler, 1961
• †Cornerocarpon Grote, 2013
  • †Cornerocarpon copiosum Grote, 2013
• †Coussapoites Pons, 1976
  • †Coussapoites veracruzianus Pons, 1976
• †Cudranioxylon Dupéron-Laudoueneix, 1980
  • †Cudranioxylon engolismense Dupéron-Laudoueneix, 1980
• †Ficofolium Peters, 1963
  • †Ficofolium weylandii Peters, 1963
• †Ficonium Ettingshausen, 1883
  • †Ficonium nitidum Paterson, 1934
  • †Ficonium silesiacum (Velenovský) Halamski & J. Kvaček, 2015
  • †Ficonium solanderi Ettingshausen, 1883
• †Milicioxylon Shukla, Mehrotra, & Guleria, 2012
  • †Milicioxylon kachchhense Shukla, Mehrotra, & Guleria, 2012
• †Moraceoipollenites Zheng, 1999
• †Moricites Krüger, 1825
• †Moroidea M. Chandler, 1957
  • †Moroidea baltica Dorofeev, 1982
  • †Moroidea caucasica Dorofeev, 1982
  • †Moroidea cretacea Knobloch & Mai, 1986
  • †Moroidea hordwellensis M. Chandler, 1961
  • †Moroidea reticulata Dorofeev, 1963
  • †Moroidea tymensis Dorofeev, 1963
• †Moroxylon Selmeier, 1993
• †Myrianthoxylon Koeniguer, 1978
  • Myrianthoxylon chaloneri Koeniguer, 1978
• †Ovicarpum M. Chandler, 1962
• †Palaeokalopanax Fotjanova, 1984
  • †Palaeokalopanax kamtschaticus Fotjanova, 1984
  • †Palaeokalopanax vollosovitschii Chelebaeva, 2005
• †Paleoficus Biswas
• †Protoficus Saporta, 1868
  • †Protoficus crenulata Saporta, 1868
  • †Protoficus crispans Langeron, 1900
  • †Protoficus dentatus Langeron, 1899
  • †Protoficus insignis Saporta, 1868
  • †Protoficus lacera Saporta, 1868
  • †Protoficus nervosa Newberry, 1883
  • †Protoficus saportae Principi, 1916
  • †Protoficus sezannensis (Watelet) Saporta, 1868
• †Soroceaxylon Franco, 2010
  • Soroceaxylon entrerriense Franco, 2010
• †Ungerites Schleiden, 1855 (syn Ficoxylon)
  • †Ungerites tropicus Schleiden, 1855
• †Welkoetoxylon Boonchai, Manchester, & Wheeler, 2015
  • †Welkoetoxylon multiseriatum Boonchai, Manchester, & Wheeler, 2015

Evolution

While the fossil record of Moraceae goes back to the late Cretaceous, molecular clock estimates suggest that the family had begun to diversify by the mid-Cretaceous, with some major clades emerging during the Tertiary period.

Distribution

Moraceae can be found throughout the world with a cosmopolitan distribution. The majority of species originate in the Old World tropics, particularly in Asia and the Pacific islands.

See also

• Barkcloth

References

1. Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III" (PDF). Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x. hdl:10654/18083. Retrieved 2013-07-06.
2. ^ Moraceae Gaudich. Plants of the World Online. Retrieved 22 April 2024.
3. Christenhusz, M. J. M.; Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3): 201–217. doi:10.11646/phytotaxa.261.3.1.
4. Judd WS, Campbell CS, Kellogg EA, Stevens PF, Donoghue MJ (2008). Plant Systematics: A Phylogenetic Approach. Sunderland, MA: Sinauer Associates, Inc. pp. 1–620. ISBN 978-0-878-93407-2.
5. Andrews, F.W. D.Sc. (1952). The Flowering Plants of the Anglo-Egyptian Sudan – Volume 2. Arbroath, Scotland: T. Buncle and Co. p. 260.
6. Thulin, M. et al. (2008). Flora of Somalia, Vol. 1–4 https://plants.jstor.org/collection/FLOS
7. Leite VG, Mansano VF, Teixeira SP (2018). "Floral Development of Moraceae species with emphasis on the perianth and androecium". Flora. 240 (Flora): 116–132. doi:10.1016/j.flora.2018.01.009.
8. TWC Staff (2018). "Morus rubra (Red Mulberry)". Wildflower.org.
9. ^ Datwyler SL, Weiblen G (2004). "On the origin of the fig: Phylogenetic relationships of Moraceae from ndhF sequences". American Journal of Botany. 91 (5): 767–777. doi:10.3732/ajb.91.5.767. PMID 21653431.
10. ^ Clement WL, Weiblen GD (2009). "Morphological evolution in the mulberry family (Moraceae)". Systematic Botany. 34 (3): 530–552. doi:10.1600/036364409789271155. S2CID 85680800.
11. ^ Sytsma KJ, Morawetz J, Pires C, Nepokroeff M, Conti E, Zjhra M, Hall JC, Chase MW (2002). "Urticalean rosids: Circumscription, rosid ancestry, and phylogenetics based on rbcL, trnL–F, and ndhF sequences" (PDF). American Journal of Botany. 89 (9): 1531–1546. doi:10.3732/ajb.89.9.1531. PMID 21665755.
12. ^ Zerega NJ, Clement WL, Datwyler SL, Weiblen GD (2005). "Biogeography and divergence times in the mulberry family (Moraceae)". Molecular Phylogenetics and Evolution. 37 (2): 402–416. CiteSeerX 10.1.1.418.1442. doi:10.1016/j.ympev.2005.07.004. PMID 16112884.
13. Gardner, E.M., Garner, M., Cowan, R., Dodsworth, S., Epitawalage, N., Arifiani, D., Sahromi, Baker, W.J., Forest, F., Maurin, O., Zerega, N.J.C., Monro, A.K. and Hipp, A. (2021), Repeated parallel losses of inflexed stamens in Moraceae: Phylogenomics and generic revision of the tribe Moreae and the reinstatement of the tribe Olmedieae (Moraceae).Taxon, 70: 946-988. https://doi.org/10.1002/tax.12526
14. Hepworth C (2018). "Moraceae – The Mulberry Family". Florida Fruit Geek.
15. "Moraceae". The International Fossil Plant Names Index. Retrieved 9 Feb 2023.
16. Zerega NJC, Clement WL, Datwyler SL, Weiblen GD (2005). "Biogeography and Divergence times in the mulberry family (Moraceae)". Molecular Phylogenetics and Evolution. 37 (2): 402–416. CiteSeerX 10.1.1.418.1442. doi:10.1016/j.ympev.2005.07.004. PMID 16112884.{{cite journal}}: CS1 maint: multiple names: authors list (link)

External links

• Moraceae in L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants.

• Moraceae
• Rosales
• Rosid families
• Extant Campanian first appearances
• Taxa named by Charles Gaudichaud-Beaupré