Hubbry Logo
FicusFicusMain
Open search
Ficus
Community hub
Ficus
logo
8 pages, 0 posts
0 subscribers
Be the first to start a discussion here.
Be the first to start a discussion here.
Ficus
Ficus
Ficus
View on Wikipedia
from Wikipedia
For the species commonly known as the "ficus tree", see Ficus benjamina. For sea snails, see Ficus (gastropod). For Monroe Ficus, see Too Close for Comfort.
"Fig tree" redirects here. For the 2009 film, see Fig Trees.

Fig trees
Temporal range: Maastrichtian–Present
Sycamore fig, Ficus sycomorus
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Rosales
Family: Moraceae
Tribe: Ficeae
Dumort.
Genus: Ficus
L.[1]
Type species
Ficus carica
L.[2]
Species

About 800, see List of Ficus species

Synonyms[1]
28 Synonyms
  • Boscheria Carruth.
  • Bosscheria de Vriese & Teijsm.
  • Caprificus Gasp.
  • Covellia Gasp.
  • Cystogyne Gasp.
  • Dammaropsis Warb.
  • Erosma Booth
  • Erythrogyne Vis.
  • Galoglychia Gasp.
  • Gonosuke Raf.
  • Macrophthalma Gasp.
  • Mastosuke Raf.
  • Necalistis Raf.
  • Oluntos Raf.
  • Perula Raf.
  • Pharmacosycea Miq.
  • Plagiostigma Zucc.
  • Pogonotrophe Miq.
  • Rephesis Raf.
  • Stilpnophyllum (Endl.) Drury
  • Sycomorphe Miq.
  • Sycomorus Gasp.
  • Synoecia Miq.
  • Tenorea Gasp.
  • Tremotis Raf.
  • Urostigma Gasp.
  • Varinga Raf.
  • Visiania Gasp.

Ficus (/ˈfaɪkəs/[3] or /ˈfiːkəs/[4][5]) 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 into the semi-warm temperate zone. The common fig (F. carica) is a temperate species native to southwest Asia and the Mediterranean region (from Afghanistan to Portugal), which has been widely cultivated from ancient times for its fruit, also referred to as figs. The fruit of most other species are also edible though they are usually of only local economic importance or eaten as bushfood. However, they are extremely important food resources for wildlife. Figs are also of considerable cultural importance throughout the tropics, both as objects of worship and for their many practical uses.

Description

[edit]
Aerial roots that may eventually provide structural support
A Ficus carica (common fig)
The stipule of Ficus religiosa. The white stipule contains a new leaf and a new stipule.
Ficus benjamina ripening fruit
Ficus watkinsiana fruit

Ficus is a pantropical genus of trees, shrubs, and vines occupying a wide variety of ecological niches; most are evergreen, but some deciduous species are found in areas outside of the tropics and to higher elevations.[6] Fig species are characterized by their unique inflorescence and distinctive pollination syndrome, which uses wasp species belonging to the family Agaonidae for pollination. Adult plants vary in size from Ficus benghalensis (the Indian banyan), a tall and speading tree with many adventitious roots which may cover a hectare (2.5 acres) or more of ground to Ficus nana of New Guinea which never exceeds one meter (forty inches) in height and width.[7]

Specific identification of many of the species can be difficult, but members of the genus Ficus are relatively easy to recognize. Many have aerial roots and a distinctive shape or habit, and their fruits distinguish them from other plants. The aerial roots according to Adrian Forsyth can hang down freely as much as 50 m (160 ft).[8] The fruit of Ficus is an inflorescence enclosed in an urn-like structure called a syconium, which is lined on the inside with the fig's tiny flowers that develop into multiple ovaries on the inside surface.[9] In essence, the fig fruit is a fleshy stem with multiple tiny flowers that fruit and coalesce.

Notably, three vegetative traits together are unique to figs. All figs present a white to yellowish latex, some in copious quantities; the twig shows paired stipules or stipular scars; the lateral veins at the base of the leaf are steep, forming a tighter angle with the midrib than the other lateral veins, a feature referred to as "triveined".

Current molecular clock estimates indicate that Ficus is a relatively ancient genus, being at least 60 million years old,[10] and possibly as old as 80 million years. The main radiation of extant species, however, may have taken place more recently, between 20 and 40 million years ago.

Some better-known species that represent the diversity of the genus include, alongside the common fig, whose fingered fig leaf is well known in art and iconography: the weeping fig (F. benjamina), a hemiepiphyte with thin, tough leaves on pendulous stalks adapted to its rain forest habitat; the rough-leaved sandpaper figs from Australia; and the creeping fig (F. pumila), a vine whose small, hard leaves form a dense carpet of foliage over rocks or garden walls.

Moreover, figs with different plant habits have undergone adaptive radiation in different biogeographic regions, leading to very high levels of alpha diversity. In the tropics, Ficus commonly is the most species-rich plant genus in a particular forest. In Asia, as many as 70 or more species can co-exist.[11] Ficus species richness declines with an increase in latitude in both hemispheres.[12][13]

A description of fig tree cultivation is set out in Ibn al-'Awwam's 12th-century agricultural work titled, Book on Agriculture.[14]

Ecology

[edit]

Figs are keystone species in many tropical forest ecosystems. Their fruit are a key resource for frugivores including fruit bats, capuchin monkeys, langurs, gibbons, and mangabeys. They are even more important for birds such as Asian barbets, pigeons, hornbills, fig-parrots, and bulbuls, which may subsist almost entirely on figs when these are plentiful. Many Lepidoptera caterpillars feed on fig leaves, for example several Euploea species (crow butterflies), the plain tiger (Danaus chrysippus), the giant swallowtail (Papilio cresphontes), the brown awl (Badamia exclamationis), and Chrysodeixis eriosoma, Choreutidae and Copromorphidae moths. The larvae of the citrus long-horned beetle (Anoplophora chinensis), for example, feed on the wood of the fig tree; the species can become a pest in fig plantations. Similarly, the sweet potato whitefly (Bemisia tabaci) is frequently found as a pest on figs grown as potted plants and can be spread through the export of these plants to other localities. For a list of other diseases common to fig trees, see List of foliage plant diseases (Moraceae).

Fig fruit and reproduction system

[edit]

See also: Fig

Many fig species are grown for their fruits, though only Ficus carica is cultivated to any extent for this purpose.[citation needed][disputed – discuss] A fig "fruit" is a type of multiple fruit known as a syconium, derived from an arrangement of many small flowers on an inverted, nearly closed receptacle. The many small flowers are unseen unless the fig is cut open.[15]

The fruit typically has a bulbous shape with a small opening (the ostiole) at the outward end that allows access to pollinators. The flowers are pollinated by very small wasps such as Pegoscapus that crawl through the opening in search of a suitable place to lay eggs. Without this pollinator service fig trees could not reproduce by seed. In turn, the flowers provide a safe haven and nourishment for the next generation of wasps. This accounts for the frequent presence of wasp larvae in the fruit, and has led to a coevolutionary relationship. Technically, a fig fruit proper would be only one of the many tiny matured, seed-bearing gynoecia found inside one fig – if you cut open a fresh fig, individual fruit will appear as fleshy "threads", each bearing a single seed inside. The genus Dorstenia, also in the fig family (Moraceae), exhibits similar tiny flowers arranged on a receptacle but in this case the receptacle is a more or less flat, open surface.[16]

Fig plants can be monoecious (hermaphrodite)[clarification needed] or gynodioecious (hermaphrodite and female).[17] Nearly half of fig species are gynodioecious, and therefore have some plants with inflorescences (syconium) with long styled pistillate flowers, and other plants with staminate flowers mixed with short styled pistillate flowers.[18] The long-styled flowers tend to prevent wasps from laying their eggs within the ovules, while the short-styled flowers are accessible for egg laying.[19]

All the native fig trees of the American continent are hermaphrodites, as well as species like Indian banyan (F. benghalensis), weeping fig (F. benjamina), Indian rubber plant (F. elastica), fiddle-leaved fig (F. lyrata), Moreton Bay fig (F. macrophylla), Chinese banyan (F. microcarpa), sacred fig (F. religiosa) and sycamore fig (F. sycomorus).[20] The common fig (Ficus carica) is a gynodioecious plant, as well as lofty fig or clown fig (F. aspera), Roxburgh fig (F. auriculata), mistletoe fig (F. deltoidea), F. pseudopalma, creeping fig (F. pumila) and related species. The hermaphrodite common figs are called "inedible figs" or "caprifigs"; in traditional culture in the Mediterranean region they were considered food for goats (Capra aegagrus). In the female fig trees, the male flower parts fail to develop; they produce the "'edible figs". Fig wasps grow in common fig caprifigs but not in the female syconiums because the female flower is too long for the wasp to successfully lay her eggs in them. Nonetheless, the wasp pollinates the flower with pollen from the caprifig it grew up in. In many situations, the wasp pollinator is unable to escape and dies within the fruit. When the wasp dies, it is broken down by enzymes (Ficain) inside the fig. Fig wasps are not known to transmit any diseases harmful to humans.

When a caprifig ripens, another caprifig must be ready to be pollinated. In temperate climes, wasps hibernate in figs, and there are distinct crops. Caprifigs have three crops per year; common figs have two.[21] The first crop (breba) is larger and juicier, and usually eaten fresh.[21] In cold climates the breba crop is often destroyed by spring frosts.[22] Some parthenocarpic cultivars of common figs do not require pollination at all, and will produce a crop of figs (albeit sterile) in the absence of caprifigs and fig wasps. According to Ziegler and Leigh, syconia "sweat" on hot days to keep the pollinating wasps from overheating. They also state that Ficus spp are the most efficient producers of sugar "of any other tree yet known".[23]

Depending on the species, each fruit can contain hundreds or even thousand of seeds.[24] Figs can be propagated by seeds, cuttings, air-layering or grafting. However, as with any plant, figs grown from seed are not necessarily genetically identical to the parent and are only propagated this way for breeding purposes.

Mutualism with the pollinating fig wasps

[edit]
Further information: Reproductive coevolution in Ficus
Ficus exasperata, fruits

The unique fig pollination system involves tiny, highly specific wasps, known as fig wasps, that enter via ostiole these subclosed inflorescences to both pollinate and lay their own eggs.[10] Each species of fig is pollinated by one or a few specialised wasp species, and therefore plantings of fig species outside of their native range results in effectively sterile individuals. For example, in Hawaii, some 60 species of figs have been introduced, but only four of the wasps that fertilize them, so only those species of figs produce viable seeds there and can become invasive species. This is an example of mutualism, in which each organism (fig plant and fig wasp) benefit each other, in this case reproductively.[15][25]

The intimate association between fig species and their wasp pollinators, along with the high incidence of a one-to-one plant-pollinator ratio have long led scientists to believe that figs and wasps are a clear example of coevolution. Morphological and reproductive behavior evidence, such as the correspondence between fig and wasp larvae maturation rates, have been cited as support for this hypothesis for many years.[26] Additionally, recent genetic and molecular dating analyses have shown a very close correspondence in the character evolution and speciation phylogenies of these two clades.[10]

According to meta-analysis of molecular data for 119 fig species 35% (41) have multiple pollinator wasp species. The real proportion is higher because not all wasp species were detected.[27] On the other hand, species of wasps pollinate multiple host fig species.[28] Molecular techniques, like microsatellite markers and mitochondrial sequence analysis, allowed a discovery of multiple genetically distinct, cryptic wasp species. Not all these cryptic species are sister taxa and thus must have experienced a host fig shift at some point.[29] These cryptic species lacked evidence of genetic introgression or backcrosses indicating limited fitness for hybrids and effective reproductive isolation and speciation.[29]

The existence of cryptic species suggests that neither the number of symbionts nor their evolutionary relationships are necessarily fixed ecologically.[29] While the morphological characteristics that facilitate the fig-wasp mutualisms are likely to be shared more fully in closer relatives, the absence of unique pairings would make it impossible to do a one-to-one tree comparison and difficult to determine cospeciation.[citation needed]

Calcium-oxalate fixation

[edit]

Several species of Ficus have been observed to sequester atmospheric CO2 as calcium oxalate in the presence of oxalotrophic bacteria and fungi, which catabolize the oxalate, which produces calcium carbonate. The calcium carbonate is precipitated throughout the tree, which also alkanalizes the surrounding soil. This process was first observed in the Iroko tree, which can sequester up to a ton of calcium carbonate in the soil over its lifespan.[30] These species are current candidates for carbon sequestration agroforestry.

Systematics

[edit]

With over 800 species, Ficus is by far the largest genus in the Moraceae, and is one of the largest genera of flowering plants currently described.[31] The species currently classified within Ficus were originally split into several genera in the mid-1800s, providing the basis for a subgeneric classification when reunited into one genus in 1867. This classification put functionally dioecious species into four subgenera based on floral characters.[32] In 1965, E. J. H. Corner reorganized the genus on the basis of breeding system, uniting these four dioecious subgenera into a single dioecious subgenus Ficus. Monoecious figs were classified within the subgenera Urostigma, Pharmacosycea and Sycomorus.[33]

This traditional classification has been called into question by recent phylogenetic studies employing genetic methods to investigate the relationships between representative members of the various sections of each subgenus.[10][32][34][35][36] Of Corner's original subgeneric divisions of the genus, only Sycomorus is supported as monophyletic in the majority of phylogenetic studies.[10][32][35] Notably, there is no clear split between dioecious and monoecious lineages.[10][32][34][35][36] One of the two sections of Pharmacosycea, a monoecious group, form a monophyletic clade basal to the rest of the genus, which includes the other section of Pharmacosycea, the rest of the monoecious species, and all of the dioecious species.[36] These remaining species are divided into two main monophyletic lineages (though the statistical support for these lineages is not as strong as for the monophyly of the more derived clades within them). One consists of all sections of Urostigma except for section Urostigma s. s.. The other includes section Urostigma s. s., subgenus Sycomorus, and the species of subgenus Ficus, though the relationships of the sections of these groups to one another are not well resolved.[10][36]

Selected species

[edit]
Main article: List of Ficus species

As of July 2025, there are 881 accepted Ficus species according to Plants of the World Online.[1]

Subgenus Ficus

[edit]

Subgenus Pharmacosycea

[edit]

Subgenus Sycidium

[edit]

Subgenus Sycomorus

[edit]

Subgenus Synoecia

[edit]

The following species[37] are typically spreading or climbing lianas:

Subgenus Urostigma

[edit]

Unplaced species

[edit]

Uses

[edit]

The wood of fig trees is often soft and the latex precludes its use for many purposes. It was used to make mummy caskets in Ancient Egypt. Certain fig species (mainly F. cotinifolia, F. insipida and F. padifolia) are traditionally used in Mesoamerica to produce papel amate (Nahuatl: āmatl). Mutuba (F. natalensis) is used to produce barkcloth in Uganda. One of the standard kbach rachana decorative elements in Cambodian architecture was inspired by the shapes of the leaves of Pou (F. religiosa). Indian banyan (F. benghalensis) and the Indian rubber plant, as well as other species, have use in herbalism.[47][citation needed] The inner bark of an unknown type of wild fig, locally known as urú, was once used by the Moré people [es] of Bolivia to produce a fibrous cloth used for clothing.[48]

Figs have figured prominently in some human cultures. There is evidence that figs, specifically the common fig (F. carica) and sycamore fig (Ficus sycomorus), were among the first plant species that were deliberately bred for agriculture in the Middle East, starting more than 11,000 years ago. Nine subfossil F. carica figs dated to about 9400–9200 BCE were found in the early Neolithic village Gilgal I (in the Jordan Valley, 13 km, or 8.1 mi, north of Jericho). These were a parthenogenetic type and thus apparently an early cultivar. This find predates the first known cultivation of grain in the Middle East by many hundreds of years.[49]

Cultivation

[edit]

Numerous species of fig are found in cultivation in domestic and office environments, including:[50]

  • F. carica, common fig – hardy to −10 °C (14 °F). Shrub or small tree which can be grown outdoors in mild temperate regions, producing substantial harvests of fruit. Many cultivars are available.
  • F. benjamina, weeping fig, ficus – hardy to 5 °C (41 °F). Widely used as an indoor plant for the home or the office. It benefits from the dry, warm atmosphere of centrally-heated interiors, and can grow to substantial heights in a favoured position. Several variegated cultivars are available.
  • F. elastica, rubber plant – hardy to 10 °C (50 °F): widely cultivated as a houseplant; several cultivars with variegated leaves
  • F. lyrata, fiddle-leaf fig – hardy to 10 °C (50 °F)
  • F. maclellandii – hardy to 5 °C (41 °F)
  • F. microcarpa, Indian laurel – hardy to 10 °C (50 °F)
  • F. pumila, creeping fig – hardy to 1 °C (34 °F)
  • F. rubiginosa, Port Jackson fig – hardy to 10 °C (50 °F)

Cultural and spiritual significance

[edit]
Further information: Fig leaf and Figs in the Bible

Fig trees have profoundly influenced culture through several religious traditions. Among the more famous species are the sacred fig tree (Pipal, bodhi, bo, or po, Ficus religiosa) and other banyan figs such as Ficus benghalensis. The oldest living plant of known planting date is a Ficus religiosa tree known as the Sri Maha Bodhi planted in the temple at Anuradhapura, Sri Lanka by King Tissa in 288 BCE. In Asia, figs are important in Buddhism and Hinduism. The Buddha is traditionally held to have found bodhi (enlightenment) while meditating for 49 days under a sacred fig.[51] The same species was Ashvattha, the "world tree" of Hinduism. The Plaksa Pra-sravana was said to be a fig tree between the roots of which the Sarasvati River sprang forth; it is usually held to be a sacred fig but more probably is Ficus virens. In Jainism, the consumption of any fruit belonging to this genus is prohibited.[52] The common fig is one of two significant trees in Islam, and there is a sura in Quran named "The Fig" or At-Tin (سوره تین). The common fig tree is first mentioned in the Bible when Adam and Eve, after gaining knowledge of their nakedness, sew fig leaves together for coverings. Throughout the Hebrew Bible, the fig tree symbolizes peace, prosperity, and divine blessing.[53] It is often paired with the grapevine as a key agricultural product of ancient Israel and is listed among the Seven Species with which the land was blessed.[53] Its sweet fruit was highly valued, and the tree appears in parables and prophetic texts, sometimes as a symbol of abundance, and at other times, when withered or destroyed, as a metaphor for judgment and desolation.[53] The fig tree was sacred in ancient Greece and Cyprus, where it was a symbol of fertility.[citation needed]

Famous fig trees

[edit]

Citations

[edit]
  1. ^ a b c "Ficus Tourn. ex L." Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 9 July 2025.
  2. ^ "Ficus L., Sp. Pl. 2: 1059 (1753)". International Plant Names Index (IPNI). Royal Botanic Gardens, Kew. 2025. Retrieved 10 June 2025.
  3. ^ "ficus". Merriam-Webster.com Dictionary. Merriam-Webster. Retrieved 2023-06-18.
  4. ^ Sunset Western Garden Book. Sunset Books. 1995. pp. 606–607. ISBN 978-0-37603-851-7.
  5. ^ "ficus". Collins English Dictionary. HarperCollins.
  6. ^ Halevy, Abraham H. (1989). Handbook of Flowering Volume 6 of CRC Handbook of Flowering. CRC Press. p. 331. ISBN 978-0-8493-3916-5. Retrieved 2009-08-25.
  7. ^ <not recorded> (2005). "Moraceae - Ficus". Flora Malesiana. 17 (part 2): 436.
  8. ^ Forsyth, Adrian (1990). Portrait of the Rainforest. Camden East: Camden House, Ontario. p. 19.
  9. ^ "Ficus: The Remarkable Genus Of Figs". Archived from the original on 2009-12-11. Retrieved 2021-05-16.
  10. ^ a b c d e f g Rønsted et al. (2005).
  11. ^ Harrison (2005).
  12. ^ van Noort & van Harten (2006).
  13. ^ Berg & Hijmann (1989).
  14. ^ Ibn al-'Awwam, Yaḥyá (1864). Le livre de l'agriculture d'Ibn-al-Awam (kitab-al-felahah) (in French). Translated by J.-J. Clement-Mullet. Paris: A. Franck. pp. 277–281 (ch. 7 - Article 25). OCLC 780050566. (pp. 277–281 (Article XXV)
  15. ^ a b "Fig Wasps". www.fs.usda.gov. Retrieved 2025-07-02.
  16. ^ Berg, Cornelis 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. 83: iii–346. ISSN 0071-5794. JSTOR 4393905.
  17. ^ Armstrong, Wayne P; Disparti, Steven (4 April 1998). "A Key to Subgroups of Dioecious* (Gynodioecious) Figs Based On Fig Wasp/Male Syconium Pollination Patterns". Wayne's Word. Archived from the original on 2012-02-02. Retrieved 2012-01-05.
  18. ^ Friis, Ib; Balslev, Henrik (2005). Plant diversity and complexity patterns: local, regional, and global dimensions. Kgl. Danske Videnskabernes Selskab. p. 472. ISBN 978-87-7304-304-2.
  19. ^ Valdeyron, Georges; Lloyd, David G. (June 1979). "Sex Differences and Flowering Phenology in the Common Fig, Ficus carica L.". Evolution. 33 (2): 673–685. doi:10.2307/2407790. JSTOR 2407790. PMID 28563939.
  20. ^ Berg & Corner (2005).
  21. ^ a b Sinha, K.K. (2003). "FIGS". Encyclopedia of Food Sciences and Nutrition. pp. 2394–2399. doi:10.1016/B0-12-227055-X/00463-6. ISBN 978-0-12-227055-0.
  22. ^ California Rare Fruit Growers, Inc. (1996): Fig Archived 2020-10-31 at the Wayback Machine. Retrieved November 1, 2008.
  23. ^ Ziegler, Christian; Leigh Jr., Egbert Giles (2002). A Magic Web. New York: Oxford University Press. p. 81. ISBN 0-19-514328-0.
  24. ^ "The Weird Sex Life of the Fig" (PDF). Ray's Figs. Retrieved 2012-01-05.
  25. ^ "The story of the fig and its wasp – Ecotone | News and Views on Ecological Science". esa.org. Retrieved 2025-07-03.
  26. ^ Machado, C. A.; Jousselin, E.; Kjellberg, F.; Compton, S. G.; Herre, E. A. (7 April 2001). "Phylogenetic relationships, historical biogeography and character evolution of fig-pollinating wasps". Proceedings of the Royal Society B: Biological Sciences. 268 (1468): 685–694. doi:10.1098/rspb.2000.1418. PMC 1088657. PMID 11321056.
  27. ^ Yang, Li-Yuan; Machado, Carlos A.; Dang, Xiao-Dong; Peng, Yan-Qiong; Yang, Da-Rong; Zhang, Da-Yong; Liao, Wan-Jin (February 2015). "The incidence and pattern of copollinator diversification in dioecious and monoecious figs". Evolution. 69 (2): 294–304. doi:10.1111/evo.12584. PMC 4328460. PMID 25495152.
  28. ^ Machado, C. A.; Robbins, N.; Gilbert, M. T. P.; Herre, E. A. (3 May 2005). "Critical review of host specificity and its coevolutionary implications in the fig/fig-wasp mutualism". Proceedings of the National Academy of Sciences. 102 (Supplement 1): 6558–6565. Bibcode:2005PNAS..102.6558M. doi:10.1073/pnas.0501840102. PMC 1131861. PMID 15851680.
  29. ^ a b c Molbo, D.; Machado, C.A.; Sevenster, J.G.; Keller, L.; Herre, E.A. (24 April 2003). "Cryptic species of fig-pollinating wasps: Implications for the evolution of the fig-wasp mutualism, sex allocation, and precision of adaptation". Proceedings of the National Academy of Sciences. 100 (10): 5867–5872. Bibcode:2003PNAS..100.5867M. doi:10.1073/pnas.0930903100. PMC 156293. PMID 12714682.
  30. ^ "From air to stone: The fig trees fighting climate change". ScienceDaily. Retrieved 2025-07-08.
  31. ^ Judd, W.S.; Campbell, C.S.; Kellogg, E.A.; Stevens, P.F.; Donoghue, M.J. (2008). Plant Systematics: A phylogenetic approach (3rd ed.). Sunderland (Massachusetts): Sinauer Associates. ISBN 978-0-87893-407-2.
  32. ^ a b c d Weiblen, G.D. (2000). "Phylogenetic relationships of functionally dioecious Ficus (Moraceae) based on ribosomal DNA sequences and morphology" (PDF). American Journal of Botany. 87 (9): 1342–1357. doi:10.2307/2656726. JSTOR 2656726. PMID 10991904. Retrieved 2018-04-22.
  33. ^ Corner, E.J.H. (1965). "Check-list of Ficus in Asia and Australasia with keys to identification". The Gardens' Bulletin Singapore. 21 (1): 1–186. Retrieved 5 Feb 2014 – via biodiversitylibrary.org.
  34. ^ a b Herre, E.; Machado, C.A.; Bermingham, E.; Nason, J.D.; Windsor, D.M.; McCafferty, S.; Van Houten, W.; Bachmann, K. (1996). "Molecular phylogenies of figs and their pollinator wasps". Journal of Biogeography. 23 (4): 521–530. Bibcode:1996JBiog..23..521H. doi:10.1111/j.1365-2699.1996.tb00014.x.
  35. ^ a b c Jousselin, E.; Rasplus, J.-Y.; Kjellberg, F. (2003). "Convergence and coevolution in a mutualism: evidence from a molecular phylogeny of Ficus". Evolution; International Journal of Organic Evolution. 57 (6): 1255–1269. doi:10.1554/02-445. PMID 12894934. S2CID 1962136.
  36. ^ a b c d Rønsted et al. (2008).
  37. ^ Berg (2003).
  38. ^ Berg (2003), p. 552.
  39. ^ Berg (2003), p. 554.
  40. ^ Berg (2003), p. 553.
  41. ^ Berg (2003), pp. 565.
  42. ^ Berg (2003), pp. 553–554.
  43. ^ Carauta & Diaz (2002), pp. 38–39.
  44. ^ a b van Noort, S.; Rasplus, J.Y. (2020). "Subsection Conosycea". Figweb: figs and fig wasps of the world. Retrieved 11 August 2019.
  45. ^ "Ficus geniculata Kurz". Plants of the World Online. Royal Botanic Gardens, Kew. 2023. Retrieved 2025-11-02.
  46. ^ Kumari, Madhu; Hemke, Jay; Chaware, Gitesh; Kinchak, Bhushan. "Ficus geniculata (Putkal): A boon". International Journal of Pharmacology and Pharmaceutical Sciences.
  47. ^ Logesh, Rajan; Vivekanandarajah Sathasivampillai, Saravanan; Varatharasan, Sujarajini; Rajan, Soundararajan; Das, Niranjan; Pandey, Jitendra; Prasad Devkota, Hari (2023-01-01). "Ficus benghalensis L. (Moraceae): A review on ethnomedicinal uses, phytochemistry and pharmacological activities". Current Research in Biotechnology. 6 100134. doi:10.1016/j.crbiot.2023.100134. ISSN 2590-2628.
  48. ^ Castedo, Luis D. Leigue (1957). El Itenez Selvaje (PDF) (in Spanish). La Paz: Ministerio de Educación. pp. 9, 16, 19, 23.
  49. ^ Kislev, Hartmann & Bar-Yosef (2006).
  50. ^ Brickell, Christopher, ed. (2008). The Royal Horticultural Society A-Z Encyclopedia of Garden Plants. United Kingdom: Dorling Kindersley. p. 448. ISBN 9781405332965.
  51. ^ "The Bodhi Tree: Uniting all Worlds". Buddhists.org. Retrieved 17 January 2020.
  52. ^ Tukol, T.K. (1980). Compendium of Jainism. Prasaranga: Karnatak University. p. 206.
  53. ^ a b c Shafer-Elliott, Cynthia (2022), Fu, Janling; Shafer-Elliott, Cynthia; Meyers, Carol (eds.), "Fruits, Nuts, Vegetables, and Legumes", T&T Clark Handbook of Food in the Hebrew Bible and Ancient Israel, T&T Clark Handbooks (1 ed.), London: T&T Clark, p. 142, ISBN 978-0-567-67982-6, retrieved 2025-07-27

General references

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Ficus

View on Grokipedia
from Grokipedia
Ficus is a of approximately 850 of flowering belonging to the mulberry family, , that are predominantly distributed across tropical and subtropical regions worldwide. These versatile exhibit a wide range of growth habits, including large trees, shrubs, woody vines (lianas), epiphytes, and hemiepiphytes, many of which produce a milky latex sap and feature simple, alternate leaves that vary from small and delicate to large and leathery. The defining characteristic of the genus is its unique , the —a fleshy, inverted receptacle that develops into the multiple-seeded —and its pollination system, which relies on an obligate mutualism with species-specific fig wasps (family ) that enter the syconium to lay eggs and pollinate the flowers in a highly specialized process. Ecologically, Ficus species are recognized as keystone plants in many ecosystems, where they provide a continuous, asynchronous fruiting cycle that sustains diverse , including birds, mammals, and , even during seasonal food shortages, thereby supporting and forest regeneration. Notable members include F. carica, the common , native to the Mediterranean and widely cultivated for its edible, nutrient-rich fruits used in food and beverages; F. elastica, the rubber plant, valued historically for used in rubber production and today as a popular ornamental ; and F. benjamina, the weeping fig, commonly grown indoors for its graceful, drooping branches. Additionally, many Ficus species hold cultural and medicinal significance in indigenous traditions, with their , leaves, and fruits employed in remedies and rituals across various societies. The genus's adaptability has led to widespread cultivation as ornamentals, though some species, like strangler figs, can become invasive in non-native habitats by overgrowing and killing host trees.

Taxonomy and Classification

Etymology and History

The genus name Ficus originates from the Latin ficus, denoting "fig" or "fig tree," a term rooted in the ancient Mediterranean where Ficus carica was one of the earliest cultivated fruits, dating back to approximately 5,000 BCE in regions like the and southwestern Asia. This plant held profound cultural and economic value in ancient civilizations, serving as a , in Greek and , and a key export across the Mediterranean basin. Carl Linnaeus formalized the genus Ficus in his seminal 1753 publication Species Plantarum, designating F. carica as the type species and encompassing a broad array of tropical and subtropical trees, shrubs, and climbers within the Moraceae family. Early post-Linnaean classifications relied on morphological traits, but the genus's complexity—due to its diverse growth forms and over 800 species—prompted major revisions in the 20th century. British botanist E.J.H. Corner advanced this work extensively from the 1950s to 1970s, producing a comprehensive check-list of Asian and Australasian species in 1958 and reorganizing the taxonomy in 1965 based on breeding systems, which consolidated the four dioecious subgenera into a single subgenus Ficus. A pivotal milestone came in 2005 with the molecular phylogenetic study by Rønsted et al., which analyzed nuclear and chloroplast DNA from 146 representative and redefined subgenera by incorporating genetic evidence, revealing deeper evolutionary relationships and challenging Corner's morphological groupings. Subsequent revisions built on this foundation; for instance, Berg and Corner's 2005 collaboration refined sectional boundaries, while a 2020 analysis by Clement et al. integrated fossil-calibrated phylogenies to clarify the genus's evolutionary history and close relatives in the Castilleae tribe. Up to 2025, taxonomic efforts have continued with phylogenomic studies on specific complexes, such as the F. erecta group in 2024, and the International Union for Conservation of Nature (IUCN) has conducted ongoing Red List assessments for numerous , evaluating threats like habitat loss to inform conservation priorities.

Phylogenetic Relationships

Ficus constitutes a monophyletic within the Moraceae, consistently supported by phylogenetic analyses of molecular data such as ndhF sequences and multi-locus datasets. These studies position Ficus as sister to the Castilleae, which includes genera like Castilla and Pourouma, highlighting its distinct evolutionary trajectory within the . Molecular phylogenetic investigations, employing markers like the (ITS) region of nuclear and the trnL-F intergenic spacer of DNA, have elucidated relationships among Ficus subgenera. These analyses generally recognize six primary subgenera—Ficus, Pharmacosycea, Sycidium, Sycomorus, Urostigma, and Synoecia—with confirmed for Sycidium, Sycomorus, and Synoecia in comprehensive sampling of over 200 species. However, subgenera such as Pharmacosycea, Urostigma, and Ficus exhibit or nesting patterns, indicating complex evolutionary histories. Urostigma, encompassing many species, emerges as one of the basal lineages in these phylogenies, particularly in reconstructions focused on the African and Australian radiations. The genus underwent an ancient radiation tied to the diversification of angiosperms, with molecular clock estimates placing the crown age of Ficus in the Eocene epoch of the period, approximately 40.6–55.9 million years ago. This timing aligns with post-Cretaceous-Paleogene boundary environmental shifts that facilitated the proliferation of tropical lineages in . Fossil evidence from the Lower further supports an early stem divergence for the genus around 73–80 million years ago. Hybridization and have played significant roles in shaping certain Ficus lineages, as revealed by genomic analyses across the . Prevalent interspecific hybridization events, detected through shared genomic blocks and admixture patterns, occurred repeatedly throughout history, complicating phylogenetic resolution in some clades. For instance, in the lineage including , evidence of hybridization with related species like F. benjamina has contributed to , while seed production via enables population persistence in certain contexts. These processes underscore the dynamic reticulate within Ficus.

Subgenera and Species Diversity

The genus Ficus encompasses approximately 850 of woody , including trees, shrubs, vines, epiphytes, and hemiepiphytes, exhibiting a distribution across , , , the , and various islands. This diversity reflects adaptations to a wide range of tropical and subtropical habitats, from rainforests to savannas. Taxonomic classification within Ficus recognizes six subgenera, as established by Berg and Corner in their 2005 revision: Pharmacosycea (monoecious, primarily Neotropical), Urostigma (monoecious, ), Ficus (gynodioecious), Sycomorus (monoecious, mostly African), Sycidium (gynodioecious, Australasian), and Synoecia (gynodioecious, Paleotropical). Subgenus Urostigma is the most species-rich, containing around 280 that dominate the genus's overall diversity, many of which are strangler figs or banyans characteristic of tropics. In contrast, subgenus Sycomorus includes fewer , approximately 15–20, centered in with extensions to the and , featuring robust trees like F. sycomorus. The remaining subgenera vary in size, with Pharmacosycea holding about 100–150 Neotropical and Sycidium around 50 Australasian ones, highlighting the genus's biogeographic partitioning. Diversity hotspots for Ficus are concentrated in and , where environmental heterogeneity supports high and . Borneo stands out as a major center, hosting over 150 —more than 18% of the —many endemic to Malesian islands due to isolation and varied . In , approximately 100–120 occur, with endemism rates exceeding 50% in regions like (23 , over 70% endemic) and the , driven by climatic stability and . These patterns underscore Ficus' role in tropical , with Southeast Asian hotspots showing up to 80 per site in some forests. Recent phylogenomic studies in the 2020s, leveraging next-generation sequencing, have refined Ficus by confirming the of subgenera like Sycomorus, Sycidium, and Synoecia, while revealing hybridization and in complexes such as F. erecta. These analyses have prompted taxonomic splits and elevations, such as recognizing new sections within Urostigma based on nuclear and markers, and the description of several new species in understudied areas like and . For instance, phylogenomic data have supported elevating certain morphologically similar groups to sectional rank, enhancing resolution of evolutionary relationships without altering the core subgeneric framework.

Morphology and Physiology

Growth Habit and Leaves

Ficus species exhibit a diverse array of growth habits, ranging from large trees and shrubs to climbers, stranglers, and woody epiphytes, all characterized by the production of milky latex in their tissues. Many species, particularly in tropical regions, grow as terrestrial trees or shrubs with a single trunk, while others function as hemiepiphytes, beginning life as epiphytes in the forest canopy before developing roots that reach the soil. Strangler figs, such as Ficus benjamina, initiate growth on host trees via seeds dispersed by birds, subsequently producing extensive aerial roots that descend to the ground, thicken into secondary trunks, and eventually envelop and often girdle the host. The leaves of Ficus are typically simple, arranged alternately on the stems (though rarely opposite or whorled), with entire margins that may be lobed in certain species like F. carica. blades vary from glabrous to hairy surfaces, often featuring pinnate or subpalmate venation and waxy spots on the abaxial side; they are supported by petioles bearing caducous stipules that form a protective sheath around emerging buds. All species produce , a milky sap exuded from wounds, which serves defensive functions against herbivores and pathogens. Some Ficus display heterophylly, where juvenile and adult leaves differ markedly in morphology to suit distinct life stages; for instance, in F. pumila, juvenile leaves are small, ovate, and adhesive for climbing support, transitioning to larger, rounded adult leaves once rooted. Physiologically, most species are , retaining leaves year-round in stable tropical climates, but patterns occur in temperate or seasonally dry environments, as seen in F. carica, which sheds leaves during winter or drought to conserve resources.

Syconium Structure and Reproduction

The syconium of Ficus represents a specialized termed a hypanthodium, characterized by an inverted, fleshy receptacle that forms a hollow, urn-like structure enclosing hundreds of minute unisexual flowers on its inner surface. This closed develops from a flattened receptacle that grows inward, creating a cavity accessible only through a small apical pore known as the ostiole, which is sealed by specialized bracts. The internal floral diversity includes male flowers, typically clustered near the ostiole, and female flowers distributed along the receptacle wall, with variations in style length that play key roles in reproductive strategies. Reproduction in Ficus occurs primarily through sexual means facilitated by the , with exhibiting either monoecious or functionally dioecious breeding systems. In monoecious , which comprise about half of the , each syconium contains both male flowers—producing —and female flowers of two types: long-styled flowers that develop into viable seeds upon , and short-styled gall flowers whose ovaries serve as sites for fig wasp oviposition and larval development. Male trees in dioecious produce syconia with male flowers and short-styled gall flowers exclusively, while female trees bear only long-styled female flowers, leading to seed production dependent on wasp . These systems ensure the of Ficus with specific pollinating wasps, where female wasps enter the syconium via the ostiole to pollinate and lay eggs. Seed development in Ficus varies by species and breeding system, with successful fertilization of long-styled flowers yielding numerous small s dispersed upon maturation and rupture. In contrast, the cultivated edible fig Ficus carica demonstrates in its (common) cultivars, where develop into fleshy, seedless fruits without or fertilization, driven by hormonal regulation that promotes expansion independently of formation. This parthenocarpic mode allows commercial production without reliance on male caprifig pollinators, though some varieties require wasp for Smyrna-type figs to set seed-bearing fruit. Anatomically, the ostiolar bracts in F. carica vary in thickness and interlocking patterns to regulate access, while internal bracts protect developing flowers, contributing to the 's role as both and precursor.

Unique Adaptations

Ficus species exhibit several specialized physiological and structural that enhance their survival in diverse environments. One prominent feature is the presence of (CaOx) crystals in their leaves, which serve dual roles in defense and mechanical support. These crystals, typically prismatic or druse forms, are deposited along the major and in tissues, often in lines parallel to the vein axes, providing rigidity to the leaf structure and contributing to overall tissue strength. In addition to structural reinforcement, CaOx crystals act as a physical deterrent against herbivores by abrading the mouthparts of chewing and forming barriers that reduce and damage from feeding. This is particularly evident in young tropical leaves, where crystal accumulation correlates with increased leaf toughness to protect vulnerable tissues during early development. The milky produced by Ficus represents another key biochemical , rich in proteolytic enzymes such as ficin, a that plays critical roles in both defense and wound response. Ficin facilitates the digestion of proteins from invading and herbivores, thereby inhibiting microbial growth and deterring insect pests at wound sites. Upon tissue damage, the rapidly coagulates to form a seal that prevents loss, blocks entry, and restores mechanical integrity to injured bark or stems, as demonstrated in like where coagulation enhances tensile strength post-injury. This rapid self-healing mechanism not only minimizes risks but also supports the plant's resilience in habitats prone to physical disturbances. In certain Ficus species, such as F. benghalensis, the strangler habit exemplifies a remarkable structural for canopy and resource . Germinating as epiphytes on host trees, these figs produce extensive that descend to the , penetrating and anchoring into the ground while expanding laterally to form a lattice around the host trunk. Over time, the thickening and lignification of these exert compressive on the host, restricting its girth expansion and vascular flow, ultimately leading to canopy takeover as the fig's branches overshadow and outcompete the host for light and nutrients. This hemiepiphytic strategy allows Ficus to exploit elevated microsites while transitioning to terrestrial dominance, thriving in understories where direct ground establishment is challenging.

Distribution and Ecology

Global Range and Habitats

The genus Ficus exhibits a distribution, spanning tropical and subtropical regions across the globe, with approximately 850 in total. Diversity is highest in and , where over 500 occur, followed by about 150 in the and roughly 100 in and . This uneven distribution reflects the genus's evolutionary history, with centers of in and . As of mid-2025, recognizes 881 accepted , highlighting ongoing taxonomic refinements. Ficus species inhabit a wide array of environments, from lowland tropical rainforests and montane forests to savannas, riverine zones, and coastal mangroves. Many thrive in wet tropical conditions but show adaptability to seasonal dry periods, with some occupying semi-arid woodlands or disturbed open habitats. Altitudinal ranges extend from to over 2,700 meters in neotropical regions, allowing occupation of diverse elevations within forest gradients. Biogeographic patterns in Ficus suggest Gondwanan origins, with ancestral lineages present in the supercontinent, contributing to disjunct distributions between Africa, South America, and Australasia through vicariance and subsequent dispersal. This ancient heritage explains the presence of related species across now-separated landmasses. In non-native regions, such as Florida, several Ficus species, including F. microcarpa, demonstrate invasive potential, establishing in urban and natural habitats and outcompeting native vegetation.

Pollination Mutualism

The pollination of Ficus relies on an mutualistic relationship with fig wasps of the family , where the wasps serve as the exclusive pollinators and the figs provide a site for wasp . This is highly specific, with each of the approximately 850 Ficus typically associated with one or a few dedicated wasp , resulting in hundreds of documented Ficus-wasp associations worldwide. The agaonid wasps, often genus-specific to their fig hosts, have coevolved with Ficus over millions of years, forming one of the most extreme examples of codiversification in plant-insect interactions. The process begins when receptive female wasps, laden with from their natal , locate a new using volatile chemical cues emitted by the . The female forces entry through the narrow ostiole at the syconium's apex, often losing her wings and antennae in the process, and deposits onto the female flowers inside. She then selectively lays eggs into some flowers using her , after which she dies within the syconium. The wasp larvae develop inside galled flowers, feeding on the floral tissue; wingless males emerge first, mate with the wingless females, and later chew exit tunnels through the syconium wall before dying. The emerging females collect from male flowers and exit to seek new figs, perpetuating the cycle. Evidence of is evident in the precise morphological adaptations between partners, such as the wasp's body size matching the ostiole dimensions to allow entry while excluding non-adapted , and synchronized phenologies aligning wasp emergence with fig receptivity. Phylogenetic studies confirm parallel diversification, with fig and wasp lineages diverging in tandem across subgenera, supporting long-term reciprocal selection. Fig wasps exhibit two pollination strategies: active and passive. In active pollination, prevalent in most Ficus species, females deliberately collect into specialized thoracic pockets and deposit it onto stigmas during oviposition, enhancing efficiency and set. Passive , found in about 30% of species (primarily Sycomorus), involves adhering incidentally to the wasp's body without deliberate placement, often requiring higher production to compensate. This mutualism incurs costs for the , as oviposited flowers develop into housing wasp offspring rather than —a form of partial —balancing production against pollinator provisioning.

Dispersal and Interactions

Ficus seeds are primarily dispersed through endozoochory, where frugivorous animals consume the ripe syconia and excrete viable seeds away from the parent plant. In tropical forests, birds such as hornbills and pigeons, bats including fruit bats like the (Pteropus giganteus), and primates such as monkeys (e.g., spider monkeys Ateles geoffroyi and proboscis monkeys Nasalis larvatus) play crucial roles as dispersers. These animals are attracted to the nutrient-rich, often brightly colored figs, which are ingested whole or partially, with seeds passing through the digestive tract unharmed due to their small size and protective coating. For instance, in Southeast Asian forests, hornbills have been observed to disperse Ficus seeds over distances of up to several kilometers, contributing to the of fig populations. Long-distance dispersal of Ficus seeds, essential for colonizing remote islands and expanding ranges, can occur via thalassochory, where seeds or entire syconia are transported by ocean currents. Recent studies have documented viable Ficus seeds floating in for extended periods, with experiments showing rates remaining high after simulated oceanic travel. This mechanism likely facilitated the colonization of isolated archipelagos, such as those in the Pacific and Indian Oceans, where Ficus species arrived post-volcanic eruptions or other disturbances without immediate animal vectors. In the case of the Krakatau Islands, oceanic drift combined with subsequent animal dispersal helped establish fig populations after the eruption. Ficus species function as keystone species in tropical ecosystems by providing a reliable, year-round food source through asynchronous fruiting across individuals and populations, sustaining during seasonal scarcities. In many tropical forests, figs support over 1,200 vertebrate species, including birds, mammals, and reptiles, which rely on them for and . This role enhances stability, as fig-dependent in turn aid in and , while the trees themselves host diverse epiphytes and . For example, in Australian and Asian rainforests, fig trees maintain frugivore populations that would otherwise decline during lean periods. Antagonistic interactions with Ficus include by , such as rats ( spp.) and mice, which consume fallen or cached seeds, reducing recruitment success. In forest understories, can remove up to 90% of accessible Ficus seeds in high-density areas, particularly for smaller-seeded . To counter herbivory, Ficus employs multiple defenses, including chemical compounds like with proteolytic enzymes that deter feeding, physical barriers such as tough leaves and crystals that abrade , and behavioral strategies like to drop infested fruits. These defenses vary by and , with pioneer Ficus like F. hispida exhibiting higher investment in anti-herbivore traits due to greater exposure.

Species Accounts

Subgenus Urostigma

The subgenus Urostigma represents the largest group within the genus Ficus, encompassing approximately 280 of primarily hemi-epiphytic strangler figs distributed across the tropics, with significant extension. These typically initiate life as epiphytes on host trees, germinating from seeds lodged in bark crevices, and develop extensive aerial adventitious roots that eventually girdle and supplant the host, forming independent trees with broad, often flat-topped crowns. The subgenus exhibits high diversity in , particularly in , where around 68 indigenous occur, many of which are endemic to islands like and , reflecting adaptive radiations in humid tropical forests. Shared traits among Urostigma species include the production of anastomosing that create supportive "root-baskets," monoecious inflorescences within syconia, and primarily by birds, which consume the fruits and excrete viable onto suitable perches. Leaves are typically spirally arranged, coriaceous, and feature a single basal waxy on the midrib underside, aiding in identification and contributing to in humid environments. These adaptations enable resilience in diverse habitats, from lowland rainforests to coastal zones, though many species face pressures from . Prominent examples include F. benghalensis (Indian banyan), renowned for its enormous crowns spanning several hectares supported by numerous prop roots, native to and often reaching heights of 20-30 meters. F. microcarpa (Chinese banyan), originating from to , is a fast-growing urban invasive with dense and smooth gray bark, frequently forming thickets in disturbed areas and used in cultivation. F. religiosa (sacred fig or bo tree), widespread in the and , features heart-shaped leaves with elongated tips and holds cultural significance, growing to 30 meters with drought-tolerant traits. These species exemplify the subgenus's ecological versatility and human associations.

Subgenus Sycomorus

The subgenus Sycomorus encompasses approximately 30 dioecious species, primarily distributed across , where they form a distinct characterized by free-standing trees rather than hemi-epiphytic habits. These species exhibit robust growth forms, often reaching heights of 20 meters or more, with dense, spreading crowns that provide substantial shade in open s and riparian zones. Unlike the strangling figs of other subgenera, Sycomorus species develop independently from seeds, establishing as terrestrial trees in , , and edge habitats. A prominent example is , the sycamore fig, which features large, leathery leaves and a smooth, yellowish bark that flakes in patches, enabling it to thrive in semi-arid conditions. Reproductive structures in the subgenus are marked by large syconia—multiple-fruited inflorescences—that develop cauliflorously on trunks and older branches, often in conspicuous clusters. Dioecy is a defining feature, with male trees bearing syconia that function as breeding sites for specialized fig wasps (genus Ceratosolen), which pollinate the flowers and lay eggs within galls; female trees, in contrast, produce seed-only syconia lacking wasp offspring, relying on animal dispersers like birds, monkeys, and elephants for seed propagation. This sexual dimorphism influences syconium size and placement, with female figs typically larger and more colorful to attract frugivores, while male figs are smaller and positioned to facilitate wasp entry and exit. Such traits have evolved convergently within the subgenus to optimize pollination mutualism and seed dispersal in fragmented African landscapes. Ecologically, Sycomorus species are keystone components of gallery forests and riverine ecosystems across , including critical areas along the Valley, where they sustain by offering asynchronous fruiting that provides reliable during dry seasons. Ficus sycomorus, in particular, supports a wide array of vertebrates and , from fruit bats to , contributing to nutrient cycling and habitat connectivity in otherwise arid environments; its presence enhances overall forest resilience and in these linear habitats. Human interactions with the subgenus date back millennia, notably with , which was cultivated for its nutritious, pear-shaped figs in since the third millennium BCE. Archaeological evidence from the Nile Valley, including root remnants from the Badarian period (circa 4000 BCE), indicates early for , timber, and shade, with practices involving gashing or puncturing the syconia to induce parthenocarpic development and improve yields. This species' edibility and adaptability made it a staple in Egyptian agriculture and diet, underscoring its enduring cultural and economic value.

Other Subgenera and Unplaced Species

Subgenus Pharmacosycea is a monophyletic group comprising approximately 70 , predominantly distributed across the , with a smaller palaeotropical contingent. It is divided into two sections: the neotropical sect. Pharmacosycea, which includes about 25 characterized by syconia with three basal bracts and often growing as tall trees in humid forests, and the palaeotropical sect. Oreosycea with around 45 adapted to diverse tropical environments. A representative example is Ficus insipida, a widespread in Amazonian riverine habitats known for its role in ecosystems. This subgenus exhibits higher in understudied regions such as Amazonia, where ongoing molecular studies continue to reveal new taxa. Subgenus Sycidium encompasses roughly 100 species, primarily as shrubs or small trees spanning from through , , and the Pacific Islands to . These are typically monoecious and feature syconia borne in leaf axils, with adaptations for by specific fig wasps in varied habitats including dry forests and montane regions. In the , related shrubby forms align with broader patterns in the , though core diversity lies in the ; examples include Ficus capreifolia in African savannas, highlighting the subgenus's ecological versatility. Phylogenetic analyses confirm its , distinguishing it from other subgenera through consistent morphological traits like scars. Smaller subgenera include and Synoecia, both centered in and . Ficus contains monoecious species, such as Ficus carica, the common , adapted to Mediterranean-like climates with edible syconia. Synoecia, with about 72 species mostly gynodioecious and often climbing or hemi-epiphytic, predominates in Malesian rainforests; recent phylogenetic work has proposed merging it with parts of Ficus (excluding subsection Ficus) into a single to reflect their close affinities, supported by genome analyses. This merger underscores the evolutionary convergence in breeding systems and habits within these groups. Several species remain unplaced or have been recently described based on molecular data, contributing to the genus's estimated total of over 850 species. For instance, Ficus sytsmae from South America was reinstated in 2025 using integrated taxonomic approaches, while Ficus naikii from central India was described in the same year, highlighting ongoing discoveries in biodiverse but understudied areas like the Amazon and Indian subcontinent. These additions emphasize patterns of undescribed diversity, particularly in tropical hotspots where molecular phylogenetics is refining classifications.

Human Interactions

Cultivation Techniques

Ficus species are primarily propagated vegetatively to maintain desirable traits, as is rare due to the dependency on specific fig wasps for and viable production in most species. Common methods include stem cuttings and air-layering, which are effective for both ornamental and fruiting varieties. For Ficus carica, hardwood cuttings taken in root readily in well-drained media under mist, while air-layering suits larger specimens of species like . Cultivation requires well-drained, organically rich soil with a neutral to slightly acidic (6.5-7.0) to prevent , particularly in tropical and subtropical climates where Ficus thrives with temperatures above 15°C (59°F) and moderate humidity. Regular watering maintains even moisture without waterlogging, and fertilization with balanced nutrients supports growth during active seasons. is essential for shape and health; for cultivation of (also known as var. retusa), cut back new growth to two or three leaves year-round to promote ramification and reduce leaf size, performed ideally in to allow recovery. Popular cultivars of F. carica for cultivation include 'Brown Turkey', valued for its cold tolerance and productivity, and 'Celeste', noted for sweet, closed-eye fruits resistant to splitting. These parthenocarpic varieties produce fruit without pollination, simplifying home and commercial growing. Pest management focuses on scale insects, common on Ficus foliage and stems, which can be controlled with horticultural oils or insecticidal soaps applied during crawler stages to avoid phytotoxicity. Integrated approaches include monitoring and encouraging natural predators like lady beetles in outdoor settings. Commercial production of F. carica centers on Mediterranean regions like , , and , where orchards yield up to 10 tons per under irrigated conditions, with global output approximately 1.3 million metric tons annually from about 299,000 s (as of 2024).

Edible and Medicinal Uses

The fruits of Ficus carica, commonly known as figs, are widely consumed fresh or dried and serve as a significant edible resource. Fresh figs contain approximately 79% water, 19% carbohydrates (primarily sugars such as glucose and ), and 0.8% protein, with a moderate content of about 2.9 grams per 100 grams, aiding and providing . Dried figs, due to , exhibit concentrated nutrients, including higher levels of (up to 14.6 grams per 100 grams) and natural sugars (around 48 grams per 100 grams), making them a popular energy-dense snack in various cuisines. Beyond F. carica, other Ficus species contribute to edible uses; for instance, in parts of , the young leaves of Ficus glomerata (synonymous with F. racemosa) are harvested and cooked as a , similar to , in traditional diets. Medicinally, Ficus species have been employed in traditional practices, particularly for dermatological and gastrointestinal issues. The milky latex from F. carica and related species contains ficin, a proteolytic enzyme that breaks down proteins, and is applied topically to treat warts, with clinical studies demonstrating its efficacy in causing wart regression within weeks without reported side effects. Ethnobotanical surveys across Asia and Africa document the use of bark decoctions from F. racemosa and F. glomerata to alleviate diarrhea, attributing this to tannins and other astringent compounds that reduce intestinal inflammation and motility. Modern research supports and expands on these traditional applications, highlighting bioactive compounds in Ficus species. Extracts from F. racemosa bark and fruits exhibit strong antioxidant activity, scavenging free radicals and reducing in vitro and in animal models, primarily due to like and . Additionally, analyses have identified sterols and in F. racemosa leaves and bark with potential antidiabetic properties, including the ability to lower blood glucose levels and improve insulin sensitivity in streptozotocin-induced diabetic rats. These findings underscore the therapeutic promise of Ficus species, though further clinical trials are needed to validate efficacy in humans.

Ornamental and Landscaping Roles

Ficus species are widely utilized as ornamental due to their attractive foliage, adaptable growth habits, and tolerance for indoor and outdoor environments. , commonly known as the weeping fig, is a popular choice for houseplants, offices, and interior , valued for its elegant form and dense, glossy dark green leaves. Similarly, , the rubber plant, is favored as an ornamental houseplant for its large, glossy evergreen leaves and broadleaf structure, often grown indoors in temperate climates to add a tropical aesthetic. These species thrive in moderate light and humidity, making them suitable for urban interiors. In , certain Ficus serve as shade trees in parks and large open spaces, providing dense canopies and structural interest. Ficus macrophylla, the fig, is planted for its grand stature, massive buttress roots, and ability to offer substantial shade in warmer climates like and . However, some pose challenges due to invasiveness; for instance, Ficus microcarpa, the Chinese banyan, has become problematic in , where it forms dense canopies that outcompete native vegetation and is listed among the world's worst invasive alien . Its aggressive root system and rapid spread can damage and ecosystems in suitable habitats, with ongoing management efforts including removal and prevention of further spread. Ficus species are also prized for bonsai and topiary applications, leveraging their resilience to pruning and distinctive aerial roots. Species like Ficus microcarpa and Ficus rubiginosa are commonly used in bonsai for their hardiness, with techniques involving regular leaf and branch pruning to maintain compact forms and encourage aerial root development through high humidity (70-100%). In topiary, Ficus benjamina tolerates severe shearing to create hedges, screens, or shaped forms, such as braided or spiraled stems, while aerial roots can be selectively pruned or trained to enhance aesthetic features without harming the plant. The global trade in potted Ficus plants contributes significantly to the ornamental , with Ficus representing a key segment due to their popularity in indoor and landscape markets.

Cultural and Conservation Aspects

Symbolic and Religious Importance

In , Ficus religiosa, known as the Bodhi tree, holds profound significance as the site where Siddhartha Gautama attained enlightenment around 528 BCE, symbolizing awakening and spiritual wisdom. This sacred fig is revered in and traditions, with descendants of the original tree propagated and venerated at sites like the in , , where pilgrims meditate beneath its branches to seek insight. In , the same , called the peepal tree, is associated with deities like and , believed to house divine spirits and used in rituals for prosperity and protection, as described in ancient texts such as the . Ficus benghalensis, the tree, features prominently in as the , or wish-fulfilling tree, embodying abundance and eternal life in epics like the , where it grants desires to the faithful. Its expansive canopy and , which allow it to live for centuries, reinforce its role as a symbol of and sustenance in Indian folklore. In modern , F. benghalensis serves as the national tree, representing unity and longevity in and environmental conservation efforts. Across African traditions, , the sycamore fig, plays a central role in rituals among groups like the of , where it is known as the mugumo tree and considered a sacred abode of , the supreme deity, used for prayers, sacrifices, and oaths to invoke rain, fertility, and community harmony. In ancient Egyptian culture, the tree symbolized protection and nourishment, linked to goddesses such as and , with its fruit offered in funerary rites to sustain the deceased in the . Figs from various Ficus species broadly symbolize due to their shape and milky sap, evoking female genitalia and lactation in Greco-Roman and Mediterranean art, as seen in depictions associated with and . Their remarkable longevity—some banyans enduring over 1,000 years—further endows them with connotations of in , from biblical parables of prosperity to Hindu tales of divine endurance. In global art and , such as paintings and ancient poetry, figs represent renewal and hidden knowledge, bridging material abundance with spiritual depth.

Notable and Famous Trees

One of the most renowned Ficus specimens is () in the Acharya Jagadish Chandra Bose Indian Botanic Garden, , . This tree, estimated to be at least 250 years old, features a vast elliptical canopy spanning approximately 190 meters in length and 145 meters in width, supported by over 3,000 aerial prop that function as additional trunks. Its expansive form covers about 1.9 hectares, making it one of the widest single trees globally and a major attraction for its demonstration of the species' characteristic growth via strangler . The Jaya Sri Maha Bodhi (Ficus religiosa) in Anuradhapura, Sri Lanka, holds the distinction as the oldest verified human-planted tree in the world. Planted in 288 BCE from a cutting of the original Bodhi tree under which Siddhartha Gautama attained enlightenment, this sacred fig is over 2,300 years old and has been meticulously preserved through cloning and protective measures. Its survival and propagation underscore the deep historical and spiritual ties of F. religiosa to Buddhist traditions. Among the largest Ficus macrophylla specimens, a standout example in boasts a trunk circumference of 29 meters, ranking it second among the nation's girthiest trees. Native to eastern , this fig exemplifies the species' potential for massive buttressed trunks and broad canopies in subtropical environments, contributing to its ecological role in rainforest canopies.

Threats and Conservation

Ficus species, predominantly native to tropical regions, are increasingly vulnerable to habitat loss driven by for , , and . Tropical forests, the primary habitats for most Ficus, have declined by approximately 50% since 1950, severely fragmenting populations and reducing suitable environments for these keystone trees. This loss disrupts the complex mutualistic relationships Ficus maintain with , including seed dispersers and pollinators. Climate change poses a profound threat to Ficus through its impacts on fig wasps, the pollinators essential for nearly all species' reproduction. Rising temperatures can exceed the thermal tolerances of these wasps during their brief life cycles, potentially leading to local extinctions and rendering Ficus unable to produce viable seeds. Studies indicate that without wasp , Ficus populations could collapse, amplifying broader ecosystem disruptions in tropical forests. Conservation assessments reveal that a notable portion of Ficus face extinction risks, with several listed as threatened on the ; for instance, the recently described Ficus naikii from is classified as Critically Endangered due to and restricted distribution. Other examples include Ficus cupulata, assessed as Endangered owing to limited locations and ongoing threats. These statuses underscore the genus's vulnerability, particularly for endemic and montane . Efforts to protect Ficus emphasize through protected areas, such as national parks in where species like Ficus insipida thrive in remnants of tropical forests, supporting and restoration initiatives. Ex situ strategies complement these by maintaining living collections and banks; for example, extensive repositories of Ficus carica cultivars in preserve for potential reintroduction. Co-conservation of fig wasps is integral, as habitat protection for Ficus inherently safeguards their pollinators, preserving the mutualistic bond critical to both. In the 2020s, genomic has advanced conservation by elucidating stress-response mechanisms in Ficus, enabling the development of resilient varieties through targeted breeding to counter and pressures. These initiatives, including phenotypic and metabolic analyses, provide foundational data for enhancing across the genus.

References

  1. https://pmc.ncbi.nlm.nih.gov/articles/PMC11645579/
  2. https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/ficus
  3. https://www.fs.usda.gov/wildflowers/pollinators/pollinator-of-the-month/fig_wasp.shtml
  4. https://www.sciencedirect.com/science/article/pii/S2351989418300969
  5. https://ethnobiomed.biomedcentral.com/articles/10.1186/s13002-020-00406-z
  6. https://edis.ifas.ufl.edu/publication/ST250
  7. https://edis.ifas.ufl.edu/publication/MG214
  8. https://www.researchgate.net/publication/346074336_The_Millenary_History_of_the_Fig_Tree_Ficus_carica_L
  9. https://www.thespruceeats.com/history-of-figs-1807598
  10. https://kosmospublishers.com/the-millenary-history-of-the-fig-tree-ficus-carica-l/
  11. https://www.etymonline.com/word/ficus
  12. https://floranorthamerica.org/Ficus
  13. https://archive.org/details/biostor-248325
  14. https://www.chicagobotanic.org/downloads/research/nyree-ronsted.pdf
  15. https://repository.naturalis.nl/pub/579346/FM1S2005017002001.pdf
  16. https://www.researchgate.net/publication/344620422_Evolution_and_classification_of_figs_Ficus_Moraceae_and_their_close_relatives_Castilleae_united_by_involucral_bracts
  17. https://www.frontiersin.org/journals/plant-science/articles/10.3389/fpls.2024.1354812/full
  18. https://www.researchgate.net/publication/387756146_Ficus_goniophylla_The_IUCN_Red_List_of_Threatened_Species
  19. https://bsapubs.onlinelibrary.wiley.com/doi/10.3732/ajb.91.5.767
  20. https://geo.cbs.umn.edu/ClementEtAl2020.pdf
  21. https://academic.oup.com/aob/article/123/1/191/5092035
  22. https://www.researchgate.net/publication/327602670_Estimating_divergence_times_and_ancestral_breeding_systems_in_Ficus_and_Moraceae
  23. https://www.nature.com/articles/s41467-021-20957-3
  24. https://www.biodiversity-science.net/EN/10.17520/biods.2018318
  25. https://onlinelibrary.wiley.com/doi/10.1002/ece3.72316
  26. https://www.inaturalist.org/taxa/50999-Ficus
  27. https://borneoficus.info/wp-content/uploads/2018/06/Berg-Corner-2005-Flora-Malesiana-Moraceae-Ficus-.pdf
  28. https://portal.cybertaxonomy.org/flora-malesiana/cdm_dataportal/taxon/d676108e-0d8e-4a5a-bd4d-3ad6387203b6
  29. https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0038432
  30. https://www.figweb.org/Ficus/Classification_of_figs/index.htm
  31. https://www.inaturalist.org/projects/moraceae-of-madagascar/journal
  32. https://academic.oup.com/bioscience/article/55/12/1053/407128
  33. https://onlinelibrary.wiley.com/doi/abs/10.1002/tax.12282
  34. https://www.treesandshrubsonline.org/articles/ficus/
  35. https://apps.cals.arizona.edu/arboretum/taxon.aspx?id=418
  36. https://hort.ifas.ufl.edu/database/documents/pdf/tree_fact_sheets/ficbena.pdf
  37. http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=112770
  38. https://naturalhistory.si.edu/research/botany/research/lianas-and-climbing-plants-neotropics/lianas-distinctive-features
  39. https://www.bbg.org/article/the_genus_ficus
  40. https://www.ias.ac.in/article/fulltext/reso/004/12/0090-0100
  41. https://pmc.ncbi.nlm.nih.gov/articles/PMC8052928/
  42. https://bsapubs.onlinelibrary.wiley.com/doi/10.2307/2656726
  43. https://nsojournals.onlinelibrary.wiley.com/doi/abs/10.1111/njb.01832
  44. https://journals.ashs.org/view/journals/hortsci/42/5/article-p1083.xml
  45. https://pmc.ncbi.nlm.nih.gov/articles/PMC5126050/
  46. https://pmc.ncbi.nlm.nih.gov/articles/PMC5813535/
  47. https://hal.science/hal-02119581v1/document
  48. http://www.scielo.sa.cr/scielo.php?script=sci_arttext&pid=S0034-77441999000100004
  49. https://www.researchgate.net/publication/319635182_Plant_Latex_Proteases_Natural_Wound_Healers
  50. https://academic.oup.com/aob/article/109/4/807/128616
  51. https://nph.onlinelibrary.wiley.com/doi/full/10.1002/ppp3.10572
  52. https://www.researchgate.net/publication/43453143_Adaptations_of_strangler_figs_to_life_in_the_rainforest_canopy
  53. https://figweb.org/Ficus/introduction/index.htm
  54. https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30001096-2
  55. https://www.researchgate.net/publication/232688038_Figs_and_the_Diversity_of_Tropical_Rainforests
  56. https://nph.onlinelibrary.wiley.com/doi/10.1111/j.1469-8137.2011.03683.x
  57. https://iforest.sisef.org/contents/?id=ifor0462-0010104
  58. https://www.nparks.gov.sg/florafaunaweb/flora/2/9/2912
  59. https://www.researchgate.net/publication/230538451_New_insight_into_the_phylogenetic_and_biogeographic_history_of_genus_Ficus_Vicariance_played_a_relatively_minor_role_compared_with_ecological_opportunity_and_dispersal
  60. https://www.iucngisd.org/gisd/speciesname/Ficus%2Bmicrocarpus
  61. https://www.pnas.org/doi/10.1073/pnas.0501840102
  62. https://pmc.ncbi.nlm.nih.gov/articles/PMC11394708/
  63. https://www.figweb.org/Fig_wasps/Species_richness/index.htm
  64. https://pmc.ncbi.nlm.nih.gov/articles/PMC3478567/
  65. https://www.frontiersin.org/journals/ecology-and-evolution/articles/10.3389/fevo.2021.685542/full
  66. https://onlinelibrary.wiley.com/doi/10.1046/j.1461-0248.2001.00209.x
  67. https://www.sciencedirect.com/science/article/pii/S0169534796100641
  68. https://www.pnas.org/doi/10.1073/pnas.1008773107
  69. https://esajournals.onlinelibrary.wiley.com/doi/abs/10.1890/04-1325
  70. https://www.sciencedirect.com/science/article/abs/pii/S1146609X22000558
  71. https://www.mdpi.com/2223-7747/13/10/1398
  72. https://www.researchgate.net/publication/229789053_The_role_of_animal_in_the_colonization_of_the_Krakatau_Islands_by_fig_tress_Ficus_species
  73. https://pmc.ncbi.nlm.nih.gov/articles/PMC4195654/
  74. https://lkcnhm.nus.edu.sg/app/uploads/2017/06/53rbz281-286.pdf
  75. https://academic.oup.com/jmammal/article/84/1/225/2373192
  76. https://pubmed.ncbi.nlm.nih.gov/30689053/
  77. https://www.nature.com/articles/s41598-025-86135-3
  78. https://pmc.ncbi.nlm.nih.gov/articles/PMC4242178/
  79. https://botany.dnp.go.th/eflora/florasubgenus.html?factsheet=Urostigma
  80. https://www.researchgate.net/publication/263494526_A_Revision_of_Ficus_Subsection_Urostigma_Moraceae
  81. https://borneoficus.info/wp-content/uploads/2018/06/Shanahan-PHD-2000-Ficus-Seed-Dispersal-Guilds.pdf
  82. https://www.gbif.org/species/144106453
  83. https://www.cabidigitallibrary.org/doi/full/10.1079/cabicompendium.24066
  84. https://www.phytologia.org/uploads/2/3/4/2/23422706/98_1_42-75riefner_ficus_microcarpa.pdf
  85. https://plants.ces.ncsu.edu/plants/ficus-microcarpa/
  86. https://www.nparks.gov.sg/florafaunaweb/flora/2/9/2915
  87. https://www.cabidigitallibrary.org/doi/full/10.1079/cabicompendium.24168
  88. https://tropical.theferns.info/viewtropical.php?id=Ficus+sycomorus
  89. https://powo.science.kew.org/taxon/853797-1
  90. https://pmc.ncbi.nlm.nih.gov/articles/PMC3367955/
  91. https://onlinelibrary.wiley.com/doi/10.1111/aje.12773
  92. https://apps.worldagroforestry.org/treedb/AFTPDFS/Ficus_sycomorus.PDF
  93. https://www.nparks.gov.sg/sbg/research/publications/gardens-bulletin-singapore/-/media/sbg/gardens-bulletin/4-4-29-22-y1977-v29-gbs-pg-191.pdf
  94. https://scholarsmine.mst.edu/cgi/viewcontent.cgi?article=1350&context=artlan_phil_facwork
  95. https://www.jstor.org/stable/4252947
  96. https://portal.cybertaxonomy.org/flora-malesiana/cdm_dataportal/taxon/f6850f9e-76e5-4383-a9c9-9b18fc0f94d2
  97. https://www.biotaxa.org/Phytotaxa/article/view/phytotaxa.313.1.1
  98. https://pmc.ncbi.nlm.nih.gov/articles/PMC11559249/
  99. https://www.researchgate.net/publication/330341978_El_genero_Ficus_subgenero_Pharmacosycea_Moraceae_en_Veracruz_Mexico
  100. http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=313023
  101. https://portal.cybertaxonomy.org/flora-malesiana/cdm_dataportal/taxon/bb20cb7d-a115-4381-bba4-a019210a67ea
  102. https://figweb.org/Ficus/Subgenus_Sycidium/Section_Sycidium/index.htm
  103. https://portal.cybertaxonomy.org/flora-malesiana/cdm_dataportal/taxon/a23c337f-4e14-4fb5-bb7a-2456a258b6ef
  104. https://pmc.ncbi.nlm.nih.gov/articles/PMC9224849/
  105. https://phytotaxa.mapress.com/pt/article/view/phytotaxa.711.3.1
  106. https://nsojournals.onlinelibrary.wiley.com/doi/abs/10.1002/njb.04788
  107. https://wfoplantlist.org/taxon/wfo-4000014727-2022-12
  108. https://www.nature.com/articles/s41598-017-09873-z
  109. https://edis.ifas.ufl.edu/publication/EP136
  110. https://plants.ces.ncsu.edu/plants/ficus-carica/
  111. https://www.kusamurabonsai.org/ficus-basics
  112. https://ipm.ucanr.edu/PMG/PESTNOTES/pn7408.html
  113. https://www.indexbox.io/blog/fig-world-market-overview-2024-2/
  114. https://www.verywellfit.com/calories-in-figs-nutrition-and-health-benefits-4109063
  115. https://foodstruct.com/food/common-fig
  116. https://prosea.prota4u.org/view.aspx?id=1660
  117. https://pubmed.ncbi.nlm.nih.gov/17472688/
  118. https://www.nature.com/articles/s41598-022-26338-0
  119. https://www.tandfonline.com/doi/full/10.3109/13880200903241861
  120. https://www.japsonline.com/admin/php/uploads/219_pdf.pdf
  121. https://pmc.ncbi.nlm.nih.gov/articles/PMC2722211/
  122. https://hgic.clemson.edu/factsheet/weeping-ficus/
  123. https://plants.ces.ncsu.edu/plants/ficus-elastica/
  124. https://edis.ifas.ufl.edu/publication/ST252
  125. https://plantselector.botanicgardens.sa.gov.au/Plants/Details/17878
  126. https://files.hawaii.gov/dbedt/erp/Doc_Library/2021-06-08-OA-FEA-Kalauhaihai-Fishpond-Management-Plan.pdf
  127. https://downloads.regulations.gov/FWS-R1-ES-2017-0050-0002/attachment_111.pdf
  128. https://www.cabidigitallibrary.org/doi/full/10.1079/cabicompendium.24130
  129. https://plants.ces.ncsu.edu/plants/ficus-microcarpa-golden-gate/
  130. https://apps.cals.arizona.edu/arboretum/taxon.aspx?id=431
  131. https://edis.ifas.ufl.edu/publication/ST251
  132. https://journals.ashs.org/view/journals/horttech/22/2/article-p169.xml
  133. https://pmc.ncbi.nlm.nih.gov/articles/PMC3249921/
  134. https://www.researchgate.net/publication/393182309_Ficus_religiosa_L_A_sacred_tree_with_medicinal_and_cultural_significance
  135. https://www.sciencedirect.com/science/article/abs/pii/S0378874111000614
  136. https://heartfulness.org/magazine/the-tree-of-life-and-fertility
  137. https://psychologynepal.com/blog/bar-and-pipal-the-two-divine-trees-in-love/
  138. https://plants.ces.ncsu.edu/plants/ficus-benghalensis/
  139. https://www.beyondforest.org/post/everything-about-the-mugumo-tree
  140. https://agroforestry.org/the-overstory/122-overstory-150-forest-and-tree-symbolism-in-folklore
  141. https://blog.metmuseum.org/cloistersgardens/2011/09/23/figs-and-fig-leaves/
  142. https://www.symbols.com/symbol/fig
  143. https://theisraelbible.com/figs-a-symbol-throughout-jewish-history/
  144. https://www.guinnessworldrecords.com/world-records/66385-tree-greatest-spread
  145. https://www.guinnessworldrecords.com/world-records/oldest-known-human-planted-tree-
  146. https://www.atlasobscura.com/places/jaya-sri-maha-bodhi-tree
  147. https://reporter.anu.edu.au/all-stories/why-tasmania-and-victoria-dominate-the-list-of-australias-largest-trees-and-why-these-majestic-giants-are-under-threat
  148. https://news.ucsc.edu/2001/02/restoration-of-tropical-forests-gets-under-way-but-scientists-still-have-a-lot-to-learn/
  149. https://www.researchgate.net/publication/236068050_Climate_warming_and_the_potential_extinction_of_fig_wasps_the_obligate_pollinators_of_figs
  150. https://pmc.ncbi.nlm.nih.gov/articles/PMC3645034/
  151. https://onlinelibrary.wiley.com/doi/full/10.1002/ece3.9311
  152. https://www.researchgate.net/publication/393794731_Ficus_naikii_Moraceae_a_new_species_from_central_India
  153. https://www.researchgate.net/publication/396447853_Ficus_cupulata_The_IUCN_Red_List_of_Threatened_Species_2024_eT239905600A239905602
  154. http://macroecointern.dk/pdf-reprints/Cottee-Jones_Biotropica_2016.pdf
  155. https://link.springer.com/article/10.1007/s11295-014-0715-3
Add your contribution
Related Hubs
User Avatar
No comments yet.