''Ficus'' ( or ) is a
genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of about 850
species
A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of woody
tree
In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only ...
s,
shrub
A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...
s,
vine
A vine is any plant with a growth habit of trailing or scandent (that is, climbing) stems, lianas, or runners. The word ''vine'' can also refer to such stems or runners themselves, for instance, when used in wicker work.Jackson; Benjamin; Da ...
s,
epiphyte
An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...
s and
hemiepiphytes in the
family
Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Moraceae
Moraceae is a family of flowering plants comprising about 48 genera and over 1100 species, 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 ...
. Collectively known as fig trees or figs, they are native throughout the
tropics
The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
with a few species extending into the semi-warm
temperate
In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
zone. The
common fig
The fig is the edible fruit of ''Ficus carica'', a species of tree or shrub in the flowering plant family Moraceae, native to the Mediterranean region, together with western and southern Asia. It has been cultivated since ancient times and is ...
(''F. carica'') is a temperate species native to southwest Asia and the
Mediterranean
The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
region (from Afghanistan to Portugal), which has been widely cultivated from ancient times for its
fruit
In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering.
Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...
, 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
Bush tucker, also called bush food, is any food native to Australia and historically eaten by Indigenous Australians and Torres Strait Islander peoples, but it can also describe any native flora, fauna, or fungi used for culinary or medicinal ...
. 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

''Ficus'' is a
pantropical
A pantropical ("all tropics") distribution is one which covers tropical regions of both the Eastern and Western hemispheres. Examples of species include caecilians, modern sirenians and the plant genera ''Acacia'' and ''Bacopa''.
''Neotropical' ...
genus of trees, shrubs, and vines occupying a wide variety of
ecological niche
In ecology, a niche is the match of a species to a specific environmental condition.
Three variants of ecological niche are described by
It describes how an organism or population responds to the distribution of Resource (biology), resources an ...
s; most are
evergreen
In botany, an evergreen is a plant which has Leaf, foliage that remains green and functional throughout the year. This contrasts with deciduous plants, which lose their foliage completely during the winter or dry season. Consisting of many diffe ...
, but some deciduous species are found in areas outside of the tropics and to higher elevations.
Fig species are characterized by their unique
inflorescence
In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...
and distinctive
pollination syndrome
Pollination syndromes are suites of flower traits that have evolved in response to natural selection imposed by different Pollination, pollen vectors, which can be abiotic (wind and water) or biotic, such as birds, bees, flies, and so forth thro ...
, which uses wasp species belonging to the family
Agaonidae
The family Agaonidae is a group of pollinating fig wasps. They spend their larval stage inside the fruits of Ficus, figs. The pollinating wasps (Agaoninae, Kradibiinae, and Tetrapusiinae) are the mutualism (biology), mutualistic partners of the ...
for pollination. Adult plants vary in size from ''
Ficus benghalensis
''Ficus benghalensis'', ''Ficus indica'', or ''Ficus audrey'' commonly known as the banyan, banyan fig and Indian banyan, is a tree native to the Indian Subcontinent. Specimens in India are among the largest trees in the world by canopy coverage. ...
'' (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
New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands by area, world's second-largest island, with an area of . Located in Melanesia in the southwestern Pacific Ocean, the island is ...
which never exceeds one meter (forty inches) in height and width.
Specific identification of many of the species can be difficult, but members of the genus ''Ficus'' are relatively easy to recognize. Many have
aerial root
Aerial roots are Root, roots growing above the ground. They are often Plant development#Adventitious structures, adventitious, i.e. formed from nonroot tissue. They are found in diverse plant species, including epiphytes such as orchids ('' ...
s 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 . The fruit of ''Ficus'' is an
inflorescence
In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...
enclosed in an urn-like structure called a
syconium
Syconium (: syconia) is the type of fruit borne by figs (genus ''Ficus''), formed by an enlarged, fleshy, hollow receptacle with multiple ovaries on the inside surface. In essence, it is really a fleshy stem with a number of flowers, so it is co ...
, which is lined on the inside with the fig's tiny flowers that develop into multiple ovaries on the inside surface. 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
Latex is an emulsion (stable dispersion) of polymer microparticles in water. Latices are found in nature, but synthetic latices are common as well.
In nature, latex is found as a wikt:milky, milky fluid, which is present in 10% of all floweri ...
, some in copious quantities; the twig shows paired
stipules
In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole). They are primarily found among dicots and rare among monocots. Stipules are considered part of the anatomy ...
—or circular scars if the stipules have fallen off; 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
The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleot ...
estimates indicate that ''Ficus'' is a relatively ancient genus, being at least 60 million years old, and possibly as old as 80 million years. The main radiation of
extant
Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to:
* Extant hereditary titles
* Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English
* Exta ...
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
The fig is the edible fruit of ''Ficus carica'', a species of tree or shrub in the flowering plant family Moraceae, native to the Mediterranean region, together with western and southern Asia. It has been cultivated since ancient times and is ...
, whose fingered
fig leaf is well known in art and
iconography
Iconography, as a branch of art history, studies the identification, description and interpretation of the content of images: the subjects depicted, the particular compositions and details used to do so, and other elements that are distinct fro ...
: the
weeping fig (''F. benjamina''), a
hemiepiphyte with thin, tough leaves on pendulous stalks adapted to its
rain forest
Rainforests are forests characterized by a closed and continuous tree Canopy (biology), canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforests can be generally classified as tropi ...
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 evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
in different
biogeographic regions, leading to very high levels of
alpha diversity
In ecology, alpha diversity (α-diversity) is the mean species diversity in a site at a local scale. The term was introduced by R. H. WhittakerWhittaker, R. H. (1960) Vegetation of the Siskiyou Mountains, Oregon and California. Ecological Monogra ...
. 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. ''Ficus''
species richness
Species richness is the number of different species represented in an community (ecology), ecological community, landscape or region. Species richness is simply a count of species, and it does not take into account the Abundance (ecology), abunda ...
declines with an increase in latitude in both hemispheres.
A description of fig tree cultivation is set out in
Ibn al-'Awwam
Ibn al-'Awwam (), also called Abu Zakariya Ibn al-Awwam (), was an Al-Andalus agriculturist who flourished at Seville (modern-day southern Spain) in the later 12th century. He wrote a lengthy handbook on agriculture entitled in Arabic '' Kitāb al ...
's 12th-century agricultural work titled, ''Book on Agriculture''.
Ecology
Figs are
keystone species
A keystone species is a species that has a disproportionately large effect on its natural environment relative to its abundance. The concept was introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in main ...
in many
tropical forest
Tropical forests are forested ecoregions with tropical climates – that is, land areas approximately bounded by the Tropic of Cancer, tropics of Cancer and Tropic of Capricorn, Capricorn, but possibly affected by other factors such as prevailing ...
ecosystem
An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
s. Their fruit are a key resource for
frugivores
A frugivore ( ) is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance ...
including
fruit bats,
capuchin monkey
The capuchin monkeys () are New World monkeys of the subfamily Cebinae. They are readily identified as the "Street organ, organ grinder" monkey, and have been used in many movies and television shows. The range of capuchin monkeys includes some t ...
s,
langurs
The Colobinae or leaf-eating monkeys are a family (biology), subfamily of the Old World monkey family that includes 61 species in 11 genus, genera, including the black-and-white colobus, the large-nosed proboscis monkey, and the gray langurs. So ...
,
gibbons Gibbons may refer to:
* Gibbon, an ape in the family Hylobatidae
* Gibbons (surname)
* Gibbons, Alberta
Gibbons is a town in central Alberta, Canada. It is located on Alberta Highway 28A, Highway 28A, northeast of Edmonton.
Gibbons is situate ...
, and
mangabeys. They are even more important for
bird
Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s such as
Asian barbets,
pigeons,
hornbill
Hornbills are birds found in tropical and subtropical Africa, Asia and Melanesia of the family Bucerotidae. They are characterized by a long, down-curved bill which is frequently brightly coloured and sometimes has a horny casque on the upper ...
s,
fig-parrots, and
bulbul
The bulbuls are members of a family, Pycnonotidae, of medium-sized passerine songbirds, which also includes greenbuls, brownbuls, leafloves, and bristlebills. The family is distributed across most of Africa and into the Middle East, tropic ...
s, which may subsist almost entirely on figs when these are plentiful. Many
Lepidoptera
Lepidoptera ( ) or lepidopterans is an order (biology), order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organ ...
caterpillar
Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths).
As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder ...
s feed on fig leaves, for example several ''
Euploea
''Euploea'' is a genus of milkweed butterflies. The species are generally dark in coloration, often quite blackish, for which reason they are commonly called crows. As usual for their subfamily, they are poisonous due to feeding on milkweeds a ...
'' species (crow butterflies), the
plain tiger (''Danaus chrysippus''), the
giant swallowtail (''Papilio cresphontes''), the
brown awl (''Badamia exclamationis''), and ''
Chrysodeixis eriosoma'',
Choreutidae and
Copromorphidae moth
Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified as suborder Heterocera, but the group is Paraphyly, paraphyletic with respect to butterflies (s ...
s. The larvae of the
citrus long-horned beetle (''Anoplophora chinensis''), for example, feed on the
wood
Wood is a structural tissue/material found as xylem in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulosic fibers that are strong in tension and embedded in a matrix of lignin t ...
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

Many fig species are grown for their fruits, though only ''
Ficus carica'' is cultivated to any extent for this purpose. A fig "fruit" is a type of
multiple fruit
Multiple fruits, also called collective fruits, are fruiting bodies formed from a cluster of flowers, the ''inflorescence''. Each flower in the inflorescence produces a fruit, but these mature into a single mass. After flowering, the mass is call ...
known as a
syconium
Syconium (: syconia) is the type of fruit borne by figs (genus ''Ficus''), formed by an enlarged, fleshy, hollow receptacle with multiple ovaries on the inside surface. In essence, it is really a fleshy stem with a number of flowers, so it is co ...
, 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.
The fruit typically has a bulbous shape with a small opening (the
ostiole
An ''ostiole'' is a small hole or opening through which algae or fungi release their mature spores.
The word is a diminutive of wikt:ostium, "ostium", "opening".
The term is also used in higher plants, for example to denote the opening of the ...
) at the outward end that allows access to
pollinator
A pollinator is an animal that moves pollen from the male anther of a flower to the female carpel, stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains.
Insects are ...
s. The flowers are pollinated by
very small wasps such as ''
Pegoscapus
''Pegoscapus'' is a genus of fig wasp native to the Americas. They range from Florida and Mexico in the north to Argentina in the south. Fig wasps have an obligate mutualism with the fig species they pollinate. ''Pegoscapus'' pollinates spec ...
'' 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
coevolution
In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well a ...
ary 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.
Fig plants can be
monoecious
Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system comparable with gynomonoecy, andromonoecy and trimonoecy, and contras ...
(
hermaphrodite
A hermaphrodite () is a sexually reproducing organism that produces both male and female gametes. Animal species in which individuals are either male or female are gonochoric, which is the opposite of hermaphroditic.
The individuals of many ...
) or
gynodioecious (hermaphrodite and female). 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. 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.
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
''Ficus sycomorus'', called the sycamore fig or the fig-mulberry (because the leaves resemble those of the mulberry), sycamore, or sycomore, is a fig species that has been cultivated since ancient times.
Etymology and naming
The specific n ...
(''F. sycomorus''). 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
goat
The goat or domestic goat (''Capra hircus'') is a species of Caprinae, goat-antelope that is mostly kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the ...
s (''Capra aegagrus''). In the female fig trees, the male flower parts fail to develop; they produce the "'edible figs".
Fig wasp
Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside fig syconia. Some are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, ...
s 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.
The first crop (
breba) is larger and juicier, and usually eaten fresh.
[ In cold climates the breba crop is often destroyed by spring frosts. Some parthenocarpic ]cultivar
A cultivar is a kind of Horticulture, cultivated plant that people have selected for desired phenotypic trait, traits and which retains those traits when Plant propagation, propagated. Methods used to propagate cultivars include division, root a ...
s of common figs do not require pollination at all, and will produce a crop of figs (albeit sterile
Sterile or sterility may refer to:
*Asepsis, a state of being free from biological contaminants
* Sterile (archaeology), a sediment deposit which contains no evidence of human activity
*Sterilization (microbiology), any process that eliminates or ...
) in the absence of caprifigs and fig wasps.
Depending on the species, each fruit can contain hundreds or even thousand of seeds. 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
The unique fig pollination
Pollination is the transfer of pollen from an anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example bees, beetles or bu ...
system involves tiny, highly specific wasps, known as fig wasp
Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside fig syconia. Some are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, ...
s, that enter via ostiole
An ''ostiole'' is a small hole or opening through which algae or fungi release their mature spores.
The word is a diminutive of wikt:ostium, "ostium", "opening".
The term is also used in higher plants, for example to denote the opening of the ...
these subclosed inflorescences to both pollinate and lay their own eggs. 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
Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
, 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
An invasive species is an introduced species that harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native spec ...
. This is an example of mutualism, in which each organism (fig plant and fig wasp
Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside fig syconia. Some are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, ...
) benefit each other, in this case reproductively.
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
In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well a ...
. 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. Additionally, recent genetic and molecular dating analyses have shown a very close correspondence in the character evolution and speciation
Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
phylogenies of these two clades.
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. On the other hand, species of wasps pollinate multiple host fig species. Molecular techniques, like microsatellite
A microsatellite is a tract of repetitive DNA in which certain Sequence motif, DNA motifs (ranging in length from one to six or more base pairs) are repeated, typically 5–50 times. Microsatellites occur at thousands of locations within an organ ...
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. These cryptic species lacked evidence of genetic introgression
Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Introg ...
or backcrosses indicating limited fitness for hybrids and effective reproductive isolation
The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, ethology, behaviors and physiology, physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensu ...
and speciation
Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
.
The existence of cryptic species suggests that neither the number of symbionts nor their evolutionary relationships are necessarily fixed ecologically. 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.
Systematics
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. 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
Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
species into four subgenera based on floral characters. 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
Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system comparable with gynomonoecy, andromonoecy and trimonoecy, and contras ...
figs were classified within the subgenera '' Urostigma'', '' Pharmacosycea'' and '' Sycomorus''.
This traditional classification has been called into question by recent phylogenetic
In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
studies employing genetic methods to investigate the relationships between representative members of the various sections of each subgenus. Of Corner's original subgeneric divisions of the genus, only ''Sycomorus'' is supported as monophyletic in the majority of phylogenetic studies. Notably, there is no clear split between dioecious and monoecious lineages. 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. 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.
Selected species
As of April 2024, there are 880 accepted ''Ficus'' species according to Plants of the World Online
Plants of the World Online (POWO) is an online taxonomic database published by the Royal Botanic Gardens, Kew.
History
Following the Convention on Biological Diversity, the Royal Botanic Gardens in Kew launched Plants of the World Online i ...
.
Subgenus ''Ficus''
* '' Ficus amplissima'' Sm. – bat fig
* '' Ficus carica'' L. – common fig
* '' Ficus daimingshanensis'' Chang
* '' Ficus deltoidea'' Jack – mistletoe fig
* '' Ficus erecta'' Thunb. – Japanese fig
* '' Ficus fulva'' Reinw. ex Blume
* '' Ficus grossularioides'' Burman f. – white-leaved fig
* '' Ficus neriifolia'' Sm.
* '' Ficus palmata'' Forssk.
* '' Ficus pandurata'' Hance
* '' Ficus simplicissima'' Lour.
* '' Ficus triloba'' Buch.-Ham. ex Voigt
Subgenus ''Pharmacosycea''
* '' Ficus crassiuscula'' Standl.
* '' Ficus gigantosyce'' Dugand
* '' Ficus insipida'' Willd.
* '' Ficus lacunata'' Kvitvik
* '' Ficus maxima'' Mill.
* '' Ficus mutabilis'' Bureau
* '' Ficus nervosa'' Heyne ex Roth
* '' Ficus pulchella'' Schott
* '' Ficus yoponensis'' Desv.
Subgenus ''Sycidium''
* '' Ficus andamanica'' Corner
* '' Ficus aspera'' G.Forst.
* '' Ficus assamica'' Miq.
* '' Ficus bojeri'' Baker
* '' Ficus capreifolia'' Delile
* '' Ficus coronata'' Spin – creek sandpaper fig
* '' Ficus fraseri'' Miq. – shiny sandpaper fig
* ''Ficus heterophylla
''Ficus heterophylla'' is a fig plant species, in the family Moraceae, which can be found in India, southern China, Indo-China and western Malesia
Malesia is a biogeographical region straddling the Equator and the boundaries of the Indomal ...
'' L.f.
* '' Ficus lateriflora'' Vahl
* '' Ficus montana'' Burm.f. – oakleaf fig
* '' Ficus opposita'' Miq. – sweet sandpaper fig
* '' Ficus phaeosyce'' K.Schum. & Lauterb.
* '' Ficus tinctoria'' G.Forst. – dye fig
* '' Ficus ulmifolia'' Lam.
* '' Ficus wassa'' Roxb.
Subgenus ''Sycomorus''
* '' Ficus auriculata'' Lour. – Roxburgh fig
* '' Ficus bernaysii'' King
* '' Ficus dammaropsis'' Diels – highland breadfruit, ''kapiak''
* '' Ficus fistulosa'' Blume
* '' Ficus hispida'' L.
* '' Ficus nota'' Merr. – ''tibig''
* '' Ficus pseudopalma'' Blanco
* '' Ficus racemosa'' L. – cluster fig
* '' Ficus septica'' Burm.f. – hauli tree
* ''Ficus sycomorus
''Ficus sycomorus'', called the sycamore fig or the fig-mulberry (because the leaves resemble those of the mulberry), sycamore, or sycomore, is a ficus, fig species that has been cultivated since ancient times.
Etymology and naming
The spec ...
'' L. – sycamore fig (Africa)
* '' Ficus variegata'' Blume
Subgenus ''Synoecia''
The following species are typically spreading or climbing liana
A liana is a long-Plant stem, stemmed Woody plant, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the Canopy (biology), canopy in search of direct sunlight. T ...
s:
* '' Ficus hederacea'' Roxb.
* '' Ficus pantoniana'' King – climbing fig
* '' Ficus pumila'' L. – creeping fig
** ''Ficus pumila'' var. ''awkeotsang'' (Makino) Corner – jelly fig
* '' Ficus punctata'' Thunb.
* '' Ficus sagittata'' J. König ex Vahl
* '' Ficus sarmentosa'' Buch.-Ham. ex Sm.
* '' Ficus trichocarpa'' Blume
* '' Ficus villosa'' Blume
Subgenus '' Urostigma''
* '' Ficus abutilifolia'' Miq.
* '' Ficus albert-smithii'' Standl.
* '' Ficus altissima'' Blume
* '' Ficus amazonica'' Miq.
* '' Ficus americana'' Aubl.
* '' Ficus aripuanensis'' Berg & Kooy
* '' Ficus arpazusa'' Carauta and Diaz – Brazil
* '' Ficus aurea'' Nutt. – Florida strangler fig
* ''Ficus benghalensis
''Ficus benghalensis'', ''Ficus indica'', or ''Ficus audrey'' commonly known as the banyan, banyan fig and Indian banyan, is a tree native to the Indian Subcontinent. Specimens in India are among the largest trees in the world by canopy coverage. ...
'' L. – Indian banyan
* '' Ficus benjamina'' L. – weeping fig
* '' Ficus bizanae'' Hutch. & Burtt-Davy
* ''Ficus blepharophylla'' Vázquez Avila
* ''Ficus broadwayi'' Urb.
* ''Ficus burtt-davyi'' Hutch.
* ''Ficus calyptroceras'' Miq.
* ''Ficus castellviana'' Dugand
* ''Ficus catappifolia'' Kunth & Bouché
* ''Ficus citrifolia'' Mill. – short-leaved fig
* ''Ficus consociata'' Carl Ludwig Blume, Bl.
* ''Ficus cordata'' Thunb.
* ''Ficus costata'' Ait.
* ''Ficus crassipes'' F.M.Bailey – round-leaved banana fig
* ''Ficus craterostoma'' Mildbr. & Burret
* ''Ficus cyathistipula'' Warb.
* ''Ficus cyclophylla'' (Miq.) Miq.
* ''Ficus dendrocida'' Kunth
* ''Ficus depressa'' Carl Ludwig Blume, Bl.
* ''Ficus destruens'' F.White
* ''Ficus drupacea'' Thunb.
* ''Ficus elastica'' Hornem. – rubber plant
* ''Ficus exasperata'' Vahl.
* ''Ficus faulkneriana'' Berg
* ''Ficus fergusonii'' (King) T.B.Worth. ex Corner
* ''Ficus glaberrima'' Blume
* ''Ficus glumosa'' Delile
* ''Ficus greiffiana'' Dugand
* ''Ficus henneana'' Miq.
* ''Ficus hirsuta'' Schott
* ''Ficus ilicina'' Miq.
* ''Ficus kurzii'' King
* ''Ficus luschnathiana'' Miq.
* ''Ficus ingens'' Miq.
* ''Ficus krukovii'' Standl.
* ''Ficus lacor'' Buch.-Ham.
* ''Ficus lapathifolia'' Miq.
* ''Ficus lauretana'' Vázquez Avila
* ''Ficus longifolia'' Schott – narrow leaf fig
* ''Ficus lutea'' Vahl
* ''Ficus lyrata'' Warb. – fiddle-leaved fig
* ''Ficus maclellandii'' King – Alii fig
* ''Ficus macrophylla'' Desf. ex Pers. – Moreton Bay fig
* ''Ficus malacocarpa'' Standl.
* ''Ficus mariae'' Berg, Emygdio & Carauta
* ''Ficus mathewsii'' Miq.
* ''Ficus matiziana'' Dugand
* ''Ficus microcarpa'' L. – Chinese banyan
* ''Ficus muelleriana'' Berg
* ''Ficus natalensis'' Hochst. – Natal fig
* ''Ficus obliqua'' G.Forst. – small-leaved fig
* ''Ficus obtusifolia'' Kunth
* ''Ficus pakkensis'' Standl.
* ''Ficus pallida'' Vahl
* ''Ficus panurensis'' Standl.
* ''Ficus pertusa'' L.f.
* ''Ficus petiolaris'' Kunth
* ''Ficus pisocarpa'' Carl Ludwig Blume, Bl.
* ''Ficus platypoda'' Cunn. – desert fig
* ''Ficus pleurocarpa'' DC. – banana fig
* ''Ficus polita'' Vahl
* ''Ficus religiosa'' L. – sacred fig
* ''Ficus roraimensis'' Berg
* ''Ficus rubiginosa'' Desf. – Port Jackson fig
* ''Ficus rumphii'' Blume
* ''Ficus salicifolia'' Vahl – willow-leaved fig
* ''Ficus sansibarica'' Warb.
* ''Ficus schippii'' Standl.
* ''Ficus schultesii'' Dugand
* ''Ficus schumacheri'' Griseb.
* ''Ficus sphenophylla'' Standl.
* ''Ficus stuhlmannii'' Warb.
* ''Ficus subcordata'' Carl Ludwig Blume, Bl.
* ''Ficus subpisocarpa'' Gagnep.
* ''Ficus subpuberula'' Corner
* ''Ficus sumatrana'' Miq.
* ''Ficus superba'' Miq.
* ''Ficus thonningii'' Blume
* ''Ficus trichopoda'' Baker
* ''Ficus trigona'' L.f.
* ''Ficus trigonata'' L.
* ''Ficus triradiata'' Corner – red-stipule fig
* ''Ficus ursina'' Standl.
* ''Ficus velutina'' Willd.
* ''Ficus verruculosa'' Warb.
* ''Ficus virens'' Aiton – white fig
* ''Ficus watkinsiana'' F.M.Bailey – Watkins' fig
Unplaced species
* ''Ficus callosa'' Willd.
* ''Ficus hebetifolia''
* '' Ficus punctata''
* ''Ficus salomonensis''
* ''Ficus tsjahela'' Burm.f.
* ''Ficus vogeliana''
Uses
The wood of fig trees is often soft and the latex
Latex is an emulsion (stable dispersion) of polymer microparticles in water. Latices are found in nature, but synthetic latices are common as well.
In nature, latex is found as a wikt:milky, milky fluid, which is present in 10% of all floweri ...
precludes its use for many purposes. It was used to make mummy Coffin, caskets in Ancient Egypt. Certain fig species (mainly Ficus cotonifolia, ''F. cotinifolia'', ''Ficus insipida, F. insipida'' and ''Ficus padifolia, F. padifolia'') are traditionally used in Mesoamerica to produce ''papel amate'' (Nahuatl: ''āmatl''). ''Mutuba'' (''Ficus natalensis, F. natalensis'') is used to produce barkcloth in Uganda. ''Pou'' (''Ficus religiosa, F. religiosa'') leaves' shape inspired one of the standard ''kbach, kbach rachana'', decorative elements in Cambodian architecture. Ficus benghalensis, Indian banyan (''F. benghalensis'') and the Indian rubber plant, as well as other species, have use in herbalism. The inner bark of an unknown type of wild fig, locally known as ''urú'', was once used by the of Bolivia to produce a fibrous cloth used for clothing.
Figs have figured prominently in some human cultures. There is evidence that figs, specifically the common fig
The fig is the edible fruit of ''Ficus carica'', a species of tree or shrub in the flowering plant family Moraceae, native to the Mediterranean region, together with western and southern Asia. It has been cultivated since ancient times and is ...
(''F. carica'') and sycamore fig (''Ficus sycomorus
''Ficus sycomorus'', called the sycamore fig or the fig-mulberry (because the leaves resemble those of the mulberry), sycamore, or sycomore, is a ficus, fig species that has been cultivated since ancient times.
Etymology and naming
The spec ...
''), 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 10th millennium BCE, 9400–9200 BCE were found in the early Neolithic village Gilgal I (in the Jordan Valley (Middle East), Jordan Valley, 13 km, or 8.1 mi, north of Tell es-Sultan, Jericho). These were a parthenogenesis, parthenogenetic type and thus apparently an early cultivar. This find predates the first known cultivation of cereal, grain in the Middle East by many hundreds of years.
Cultivation
Numerous species of fig are found in cultivation in domestic and office environments, including:
* ''Ficus carica, F. carica'', common fig – hardy to . Shrub or small tree which can be grown outdoors in mild temperate regions, producing substantial harvests of fruit. Many cultivars are available.
* ''Ficus benjamina, F. benjamina'', weeping fig, ficus – hardy to . 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.
* ''Ficus elastica, F. elastica'', rubber plant – hardy to : widely cultivated as a houseplant; several cultivars with variegated leaves
* ''Ficus lyrata, F. lyrata'', fiddle-leaf fig – hardy to
* ''Ficus maclellandii, F. maclellandii'' – hardy to
* ''Ficus microcarpa, F. microcarpa'', Indian laurel – hardy to
* ''Ficus pumila, F. pumila'', creeping fig – hardy to
* ''Ficus rubiginosa, F. rubiginosa'', Port Jackson fig – hardy to
Cultural and spiritual significance
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. The same species was ''Ashvattha'', the "world tree" of Hinduism. The ''Sacred fig, 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. 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 cited in the Bible: Adam and Eve cover their nakedness with fig leaves. The fig fruit is one of the Seven Species, traditional crops of Israel, and is included in the list of food found in the Promised Land. The fig tree was sacred in ancient Ancient Greece, Greece and Cyprus, where it was a symbol of fertility.
Famous fig trees
* ''Ashvattha'' – the world tree of Hinduism, held to be a supernatural ''F. religiosa''
* Bodhi tree – a ''F. religiosa''
* Charybdis Fig Tree of Homer's ''Odyssey'', presumably a ''F. carica''
* Curtain Fig Tree – a ''F. virens''
* Ficus Ruminalis – a ''F. carica''
* ''Plaksa'' – another supernatural fig in Hinduism; usually identified as ''F. religiosa'' but is probably ''F. virens''
* Santa Barbara's Moreton Bay Fig Tree – a ''F. macrophylla''
* Sri Maha Bodhi – another ''F. religiosa'', planted in 288 BCE, the oldest human-planted tree on record
* ''The Barren Fig Tree'' – Matthew 21:19 of the Christian Bible, Jesus Cursing of the fig tree, put a curse on the tree and used this as an example for believers of the promise of the power faith in the only true God. ''
* The Great Banyan – a ''F. benghalensis'', a clonal colony and once the largest organism known
* Vidurashwatha – "Vidura's Sacred Fig Tree", a village in India named after a famous ''F. religiosa'' that until recently stood there
* Wonderboom (tree), Wonderboom – the largest fig tree in Pretoria, South Africa, which has grown very large, through self-layering (horticulture), layering (limbs laying in the ground take root).
Citations
General references
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Supporting Online Material
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External links
��Major reference site for the genus ''Ficus''
World checklist of Ficus species from the Catalogue of Life
845 species supplied by M. Hassler's World Plants.
��Multi-award-winning documentary
BBC: Fig fossil clue to early farming
Video
How the fig tree strangles other plants for survival in the rainforest
{{Authority control
Ficus,
Moraceae genera
Taxa named by Carl Linnaeus