View source for Moraceae

{{Short description|Family of flowering plants}}
{{Automatic taxobox
| fossil_range = [[Cretaceous]] – Recent, {{fossil range|80|0}}
| image = Koeh-174.jpg
| image_caption = Panama rubber tree (''[[Castilla elastica]]'')
| taxon = Moraceae
| authority = [[Charles Gaudichaud-Beaupré|Gaudich.]] (1835) nom. cons.<ref name=APGIII2009>{{Cite journal |last=Angiosperm Phylogeny Group |year=2009 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III |journal=[[Botanical Journal of the Linnean Society]] |volume=161 |issue=2 |pages=105–121 |url=http://onlinelibrary.wiley.com/doi/10.1046/j.1095-8339.2003.t01-1-00158.x/pdf | format= PDF |access-date=2013-07-06 |doi=10.1111/j.1095-8339.2009.00996.x |doi-access=free |hdl=10654/18083 |hdl-access=free |url-access=subscription }}</ref><ref name = powo/>
| subdivision_ranks = Genera
| subdivision = 48; see text
|subdivision_ref = <ref name = powo>[https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30000043-2 Moraceae Gaudich.] ''[[Plants of the World Online]]''. Retrieved 22 April 2024.</ref>
}}

'''Moraceae''' is a family of [[flowering plant]]s comprising about 48 genera and over 1100 species,{{R|powo}}<ref name="Christenhusz-Byng2016">{{cite journal |author1=Christenhusz, M. J. M. |author2=Byng, J. W. | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = [[Phytotaxa]] | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 | doi-access = free |bibcode=2016Phytx.261..201C }}</ref> and is commonly known as the '''mulberry''' or '''fig family'''. Most are widespread in tropical and subtropical regions, less so in temperate climates; however, their distribution is cosmopolitan overall. The only common characteristics within the family are the presence of latex-producing glands in the leaves and stems, and milky sap in the soft tissues; but generally useful field characters include two [[Gynoecium|carpels]] sometimes with one reduced, compound inconspicuous flowers, and compound fruits.<ref>{{cite book |vauthors=Judd WS, Campbell CS, Kellogg EA, Stevens PF, Donoghue MJ | year = 2008 | title = Plant Systematics: A Phylogenetic Approach | publisher = Sinauer Associates, Inc. | location = Sunderland, MA | pages = 1–620 | isbn = 978-0-878-93407-2}}</ref> The family includes well-known plants such as the common fig, [[breadfruit]], [[jackfruit]] and mulberry.{{cn|date=June 2025}} The 'flowers' of Moraceae are often [[pseudanthium|pseudanthia]] (reduced [[inflorescence]]s).{{cn|date=June 2025}}

[[File:Ficus retusa, Moraceae, Bagh-e-Jinnah.jpg|thumb|''[[Ficus retusa]]'' (Moraceae) in [[Bagh-e-Jinnah, Lahore]]]]

==Description==
===Overall===
The family varies from colossal trees like the Indian Banyan (''[[Ficus benghalensis]]'') which can cover {{cvt|2|ha}} of ground, to ''[[Dorstenia barnimiana]]'' which is a small stemless, bulbous [[succulent]] 2–5&nbsp;cm in diameter that produces a single [[peltate]] leaf on a 4–15&nbsp;cm [[Petiole (botany)|petiole]]. These two species have an approximately one billion fold difference in mass.<ref>{{cite book | last= Andrews | first=  F.W. D.Sc. | date= 1952 | title= The Flowering Plants of the Anglo-Egyptian Sudan – Volume 2  | location=  Arbroath, Scotland | publisher=  T. Buncle and Co.| page= 260}}</ref><ref>Thulin, M. et al. (2008). Flora of Somalia, Vol. 1–4 https://plants.jstor.org/collection/FLOS</ref>

===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 (plant)|Castilla]]'', ''[[Dorstenia]]'', ''[[Ficus]]'', and ''[[Mesogyne]]''.<ref name="Leite, Mansano, Teixeira">{{cite journal |vauthors=Leite VG, Mansano VF, Teixeira SP | year = 2018 | title = Floral Development of Moraceae species with emphasis on the perianth and androecium | journal = Flora| volume = 240 | issue = Flora | pages = 116–132 | doi = 10.1016/j.flora.2018.01.009| doi-access =  | bibcode = 2018FMDFE.240..116L }}</ref>

===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.{{citation needed|date=August 2022}} 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.<ref name = "TWC Staff">{{cite web | author = TWC Staff | year = 2018 | title = Morus rubra (Red Mulberry) | website = Wildflower.org | url = https://www.wildflower.org/plants/result.php?id_plant=moru2}}</ref>

===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''.<ref name="Datwyler"/><ref name="Clement">{{cite journal |vauthors=Clement WL, Weiblen GD | year = 2009 | title = Morphological evolution in the mulberry family (Moraceae) | journal = [[Systematic Botany]] | volume = 34 | issue = 3 | pages = 530–552 | doi = 10.1600/036364409789271155| bibcode = 2009SysBo..34..530C | s2cid = 85680800 }}</ref>

==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.<ref name="Sytsma">{{cite journal |vauthors=Sytsma KJ, Morawetz J, Pires C, Nepokroeff M, Conti E, Zjhra M, Hall JC, Chase MW | year = 2002 | title = Urticalean rosids: Circumscription, rosid ancestry, and phylogenetics based on ''rbcL'', ''trnL''–''F'', and ''ndhF'' sequences | journal = [[American Journal of Botany]] | volume = 89 | issue = 9 | pages = 1531–1546 | url = http://www.bricol.net/research/CICphylog/CICfinalSTUFF/Urtical-rosids.pdf | doi = 10.3732/ajb.89.9.1531 | pmid = 21665755| doi-access = free }}</ref>

[[Dioecy]] (having individuals with separate sexes) appears to be the primitive state in Moraceae.<ref name="Datwyler">{{cite journal |vauthors=Datwyler SL, Weiblen G | year = 2004 | title = On the origin of the fig: Phylogenetic relationships of Moraceae from ''ndhF'' sequences | journal = [[American Journal of Botany]] | volume = 91 | issue = 5 | pages = 767–777 | doi = 10.3732/ajb.91.5.767 | pmid = 21653431| doi-access =  | bibcode = 2004AmJB...91..767D }}</ref> [[Plant sexuality#Individual plant sexuality|Monoecy]] has evolved independently at least four times within the family.

===Phylogeny===
Modern [[molecular phylogenetics]] suggest these relationships:<ref name="Sytsma"/><ref name="Datwyler"/><ref name="Clement"/><ref name="Zerega">{{cite journal |vauthors=Zerega NJ, Clement WL, Datwyler SL, Weiblen GD | year = 2005 | title = Biogeography and divergence times in the mulberry family (Moraceae) | journal = [[Molecular Phylogenetics and Evolution]] | volume = 37 | issue = 2 | pages = 402–416| doi = 10.1016/j.ympev.2005.07.004 | pmid = 16112884| bibcode = 2005MolPE..37..402Z | citeseerx = 10.1.1.418.1442 }}</ref>
{{Clade| style=line-height:75%;
|1={{clade
  |1=[[Urticaceae]]&nbsp;([[Outgroup (cladistics)|outgroup]])
  |label2=Moraceae
  |2={{clade
    |1={{clade
      |label1=[[Artocarpeae]] <!-- Missing: Hullettia and Treculia. -->
      |1={{clade
        |1={{clade
          |1=''[[Batocarpus]]''
          |2=''[[Clarisia]]''
           }}
        |2={{clade
          |1=''[[Artocarpus]]''
          |2={{clade
            |1=''[[Parartocarpus]]''
            |2=''[[Prainea]]''
             }}
           }}
         }}
      |label2=[[Moreae]]
      |2={{clade
        |1=''[[Sorocea]]''
        |2={{clade
          |1=''[[Bagassa]]''
          |2={{clade
            |1={{clade
              |1=''[[Milicia (plant)|Milicia]]''
              |2=''[[Streblus]]''
               }}
            |2={{clade
              |1=''[[Morus (plant)|Morus]]''
              |2=''[[Trophis]]''
               }}
             }}
           }}
         }}
       }}
    |2={{clade
      |label1=[[Maclureae]]
      |1=''[[Maclura]]''
      |2={{clade
        |label1=[[Dorstenieae]] <!-- Missing: Bosqueiopsis and Scyphosyce. -->
        |1={{clade
          |1=''[[Fatoua]]''
          |2={{clade
            |1={{clade
              |1=''[[Broussonetia]]''
              |2=''[[Malaisia]]''
               }}
            |2={{clade
              |1={{clade
                |1=''[[Bleekrodea]]''
                |2=''[[Sloetia]]''
                 }}
              |2=''[[Trilepisium]]''
              |3={{clade
                |1={{clade
                  |1=''[[Utsetela]]''
                  |2=''[[Dorstenia]]''
                   }}
                |2={{clade
                  |1=''[[Brosimum]]''
                  |2=''[[Trymatococcus]]''
                  |3=''[[Helianthostylis]]''
                   }}
                 }}
               }}
             }}
           }}
        |2={{clade
          |label1=[[Ficeae]]
          |1=''[[Ficus]]''
          |label2=[[Castilleae]]
          |2={{clade
            |label1=Antiaropsineae
            |1={{clade
              |1=''[[Sparattosyce]]''
              |2=''[[Antiaropsis]]''
               }}
            |label2=Castillineae
            |2={{clade
              |1={{clade
                |1=''[[Antiaris]]''
                |2=''[[Mesogyne]]''
                 }}
              |2={{clade
                |1=''[[Naucleopsis]]''
                |2={{clade
                  |1=''[[Perebea]]''
                  |2=''[[Pseudolmedia]]''
                  |3={{clade
                    |1=''[[Maquira]]''
                    |2={{clade
                      |1=''[[Helicostylis]]''
                      |2={{clade
                        |1=''[[Poulsenia]]''
                        |2=''[[Castilla (plant)|Castilla]]''
                         }}
                       }}
                     }}
                   }}
                 }}
               }}
             }}
           }}
         }}
       }}
     }}
   }}
}}

===Tribes and genera===
Moraceae comprises 48 genera<ref name = powo/> in seven tribes.<ref>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</ref>

{{div col}}
*'''[[Artocarpeae]]''' {{small|Lam. & DC. 1806}}
** ''[[Artocarpus]]'' {{small|J.R.Forst. & G.Forst.}} (73 spp.)
** ''[[Batocarpus]]'' {{small|H.Karst.}} (3 spp.)
** ''[[Clarisia]]'' {{small|Ruiz & Pav.}} (4 spp.)
* '''[[Chlorophoreae]]''' (syn. '''[[Maclureae]]''' {{small|W.L. Clement & Weiblen 2009}})
** ''[[Maclura]]'' {{small|Nutt.}} (13 spp.)
* '''Parartocarpeae'''
** ''[[Hullettia]]'' {{small|King ex Hook.f.}} (2 spp.)
** ''[[Parartocarpus]]'' {{small|Baill.}} (2 spp.)
** ''[[Pseudostreblus]]'' {{small|Bureau}} (1 sp.)
*'''[[Olmedieae]]''' {{small|Trécul 1847}} (syn. Castilleae)
** ''[[Antiaris]]'' {{small|Lesch.}} (1 sp.)
** ''[[Antiaropsis]]'' {{small|K.Schum.}} (2 spp.)
** ''[[Castilla (plant)|Castilla]]'' {{small|Cerv.}} (3 spp.)
** ''[[Helicostylis]]'' {{small|Trécul}} (8 spp.)
** ''[[Maquira]]'' {{small|Aubl.}} (4 spp.)<!-- Revista Brasileira de Psiquiatria 28(4): 277 -->
** ''[[Mesogyne]]'' {{small|Engl.}} (1 sp.)
** ''[[Naucleopsis]]'' {{small|Miq.}} (25 spp.)
** ''[[Olmedia]]'' {{small|Ruiz & Pav.}} (1 sp.)
** ''[[Perebea]]'' {{small|Aubl.}} (10 spp.)
** ''[[Poulsenia]]'' {{small|Eggers}} (1 sp.)
** ''[[Pseudolmedia]]'' {{small|Trécul}} (11 spp.)
** ''[[Sparattosyce]]'' {{small|Bur.}} (2 spp.)
** ''[[Streblus]]'' {{small|Lour.}} (5 spp.)
*'''[[Dorstenieae]]''' {{small|Dumort. 1830}}
** ''[[Bleekrodea]]'' {{small|Blume}} (3 spp.)
** ''[[Bosqueiopsis]]'' {{small|De Wild. & T.Durand}} (1 sp.)
** ''[[Brosimum]]'' {{small|Sw.}} (19 spp.)
** ''[[Broussonetia]]'' {{small|L’Hér. ex Vent.}} (4 spp.)
** ''[[Dorstenia]]'' {{small|L.}} (122 spp.)
** ''[[Fatoua]]'' {{small|Gaudich.}} (3 spp.)
** ''[[Malaisia]]'' {{small|Blanco 1837}} (1 sp.)
** ''[[Scyphosyce]]'' {{small|Baill.}} (3 spp.)
** ''[[Sloetia]]'' {{small|Teijsm. & Binn. ex Kurz}} (1 sp.)
** ''[[Sloetiopsis]]'' {{small|Engl.}} (1 sp.)
** ''[[Trilepisium]]'' {{small|Thouars}} (2 sp.)
** ''[[Utsetela]]'' {{small|Pellegr.}} (2 sp.)
*'''[[Ficeae]]''' {{small|Gaudich. 1830}}
**''[[Ficus]]'' {{small|L.}} (880 spp.)
*'''[[Moreae]]''' {{small|Dumort. 1829}}
** ''[[Afromorus]]'' {{small|E.M.Gardner}}
** ''[[Ampalis]]'' {{small|Bojer}} (2 spp.)
** ''[[Bagassa]]'' {{small|Aubl.}} (1 sp.)
** ''[[Maillardia]]'' {{small|Frapp. ex Duch.}} (2 spp.)
** ''[[Milicia (plant)|Milicia]]'' {{small|Sim}} (2 spp.)
** ''[[Morus (plant)|Morus]]'' {{small|L.}} (17 spp.)
** ''[[Paratrophis]]'' {{small|Blume}} (12 spp.)
** ''[[Sorocea]]'' {{small|A.St.-Hil.}} (22 spp.)
** ''[[Taxotrophis]]'' {{small|Blume}} (6 spp.)
** ''[[Trophis]]'' {{small|P.Browne}} (5 spp.)
{{div col end}}

Other genera accepted by [[Plants of the World Online]] {{As of|2024|April|lc=yes}}:
{{div col}}
* ''[[Allaeanthus]]'' {{small|Thwaites}} (4 spp.)
* ''[[Calaunia]]'' {{small|Gudzins.}} (1 sp.)
* ''[[Hijmania]]'' {{small|M.D.M.Vianna}} (4 spp.)
* ''[[Prainea]]'' {{small|King ex Hook.f.}} (2 spp.)
* ''[[Treculia]]'' {{small|Decne. ex Trécul}} (5 spp.)
{{div col end}}

===Fossil genera and species===
In addition to the living species, a number of fossil genera have been ascribed to the family:<ref name="IFPNI2023">{{cite web |url=http://ifpni.org/supragenus.htm?id=D69354B1-79FB-4BBE-B97C-4A3CC30B6D30 |website=The International Fossil Plant Names Index |title=Moraceae |accessdate=9 Feb 2023}}</ref>
{{div col}}
* {{extinct}}''[[Aginoxylon]]'' {{small|Dupéron, 1977}}
*''[[Aginoxylon moroides]]'' {{small|Dupéron, 1977}}
* †''[[Artocarpidium]]'' {{small|Unger, 1850}}
* †''[[Artocarpoides]]'' {{small|Saporta, 1865}}
* †''[[Arthmiocarpus]]'' {{small|Delevoryas, 1964}}
* †''[[Artocarpoxylon]]'' {{small|Prakash & Lalitha, 1978}}
* †''[[Becktonia]]'' {{small|M. Chandler, 1961}}
** †''[[Becktonia hantonensis]]'' {{small|M. Chandler, 1961}}
* †''[[Cornerocarpon]]'' {{small|Grote, 2013}}
** †''[[Cornerocarpon copiosum]]'' {{small|Grote, 2013}}
* †''[[Coussapoites]]'' {{small|Pons, 1976}}
** †''[[Coussapoites veracruzianus]]'' {{small|Pons, 1976}}
* †''[[Cudranioxylon]]'' {{small|Dupéron-Laudoueneix, 1980}}
** †''[[Cudranioxylon engolismense]]'' {{small|Dupéron-Laudoueneix, 1980}}
* †''[[Ficofolium]]''  {{small|Peters, 1963}}
** †''[[Ficofolium weylandii]]''  {{small|Peters, 1963}}
* †''[[Ficonium]]'' {{small|Ettingshausen, 1883}}
** †''[[Ficonium nitidum]]'' {{small|Paterson, 1934}}
** †''[[Ficonium silesiacum]]'' {{small|(Velenovský) Halamski & J. Kvaček, 2015}}
** †''[[Ficonium solanderi]]'' {{small|Ettingshausen, 1883}}
* †''[[Milicioxylon]]'' {{small|Shukla, Mehrotra, & Guleria, 2012}}
** †''[[Milicioxylon kachchhense]]'' {{small|Shukla, Mehrotra, & Guleria, 2012}}
* †''[[Moraceoipollenites]]'' {{small|Zheng, 1999}}
* †''[[Moricites]]'' {{small|Krüger, 1825}}
* †''[[Moroidea]]'' {{small|M. Chandler, 1957}}
** †''[[Moroidea baltica]]'' {{small|Dorofeev, 1982}}
** †''[[Moroidea caucasica]]'' {{small|Dorofeev, 1982}}
** †''[[Moroidea cretacea]]'' {{small|Knobloch & Mai, 1986}}
** †''[[Moroidea hordwellensis]]'' {{small|M. Chandler, 1961}}
** †''[[Moroidea reticulata]]'' {{small|Dorofeev, 1963}}
** †''[[Moroidea tymensis]]'' {{small|Dorofeev, 1963}}
* †''[[Moroxylon]]'' {{small|Selmeier, 1993}}
* †''[[Myrianthoxylon]]'' {{small|Koeniguer, 1978}}
**''[[Myrianthoxylon chaloneri]]'' {{small|Koeniguer, 1978}}
* †''[[Ovicarpum]]'' {{small|M. Chandler, 1962}}
* †''[[Palaeokalopanax]]'' {{small|Fotjanova, 1984}}
**†''[[Palaeokalopanax kamtschaticus]]'' {{small|Fotjanova, 1984}}
**†''[[Palaeokalopanax vollosovitschii]]'' {{small|Chelebaeva, 2005}}
* †''[[Paleoficus]]'' {{small|Biswas}}
* †''[[Protoficus]]'' {{small|Saporta, 1868}}
**†''[[Protoficus crenulata]]'' {{small|Saporta, 1868}}
**†''[[Protoficus crispans]]'' {{small|Langeron, 1900}}
**†''[[Protoficus dentatus]]'' {{small|Langeron, 1899}}
**†''[[Protoficus insignis]]'' {{small|Saporta, 1868}}
**†''[[Protoficus lacera]]'' {{small|Saporta, 1868}}
**†''[[Protoficus nervosa]]'' {{small|Newberry, 1883}}
**†''[[Protoficus saportae]]'' {{small|Principi, 1916}}
**†''[[Protoficus sezannensis]]'' {{small|(Watelet) Saporta, 1868}}
* †''[[Soroceaxylon]]'' {{small|Franco, 2010}}
**''[[Soroceaxylon entrerriense]]'' {{small|Franco, 2010}}
* †''[[Ungerites]]'' {{small|Schleiden, 1855}} (syn ''Ficoxylon'')
**†''[[Ungerites tropicus]]'' {{small|Schleiden, 1855}}
* †''[[Welkoetoxylon]]'' {{small|Boonchai, Manchester, & Wheeler, 2015}}
**†''[[Welkoetoxylon multiseriatum]]'' {{small|Boonchai, Manchester, & Wheeler, 2015}} 
{{div col end}}

==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)|Tertiary]] period.<ref name="Zerega"/>

==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.<ref name = "Zerega et al">{{cite journal | author = Zerega NJC, Clement WL, Datwyler SL, Weiblen GD | year = 2005 | title = Biogeography and Divergence times in the mulberry family (Moraceae) | journal =  Molecular Phylogenetics and Evolution| volume = 37 | issue = 2 | pages = 402–416 | doi = 10.1016/j.ympev.2005.07.004 | pmid = 16112884 | bibcode = 2005MolPE..37..402Z | citeseerx = 10.1.1.418.1442 }}</ref>

==See also==
* [[Barkcloth]]

==References==
{{Reflist}}

== External links ==
{{Wikispecies}}
{{Commons category}}
* [http://delta-intkey.com/angio/www/moraceae.htm Moraceae] in [http://delta-intkey.com/angio/ L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants.]

{{Moraceae genera}}
{{Angiosperm families}}
{{Taxonbar|from=Q156579}}
{{Authority control}}

[[Category:Moraceae| ]]
[[Category:Rosales]]
[[Category:Rosid families]]
[[Category:Extant Campanian first appearances]]
[[Category:Taxa named by Charles Gaudichaud-Beaupré]]