Ficus

Fig trees
Sycamore Fig, Ficus sycomorus
Scientific classification
Kingdom: Plantae Division: Magnoliophyta Class: Magnoliopsida Subclass: Rosidae (unranked): Eurosids I Order: Rosales Family: Moraceae Genus: Ficus L.
Species
About 800, see text

Ficus is a genus of about 850 species of woody trees, shrubs, vines, epiphytes, and hemi-epiphytes 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 so-called Common Fig (F. carica) is a temperate species from the Middle East and eastern Europe (mostly Ukraine), 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 paramount cultural importance throughout the tropics, both as objects of worship and for their many practical uses.

Contents 1 Description 2 Ecology and uses 3 Cultural and spiritual significance 4 Fig pollination and fig fruit 5 Selected species 6 List of famous fig trees 7 See also 8 Footnotes 9 References 10 External links

Description

Figs occupy a wide variety of ecological niches. Take, for example, the Common Fig, a small temperate deciduous tree whose fingered fig leaf is well-known in art and iconography; or the Weeping Fig (F. benjamina) a hemi-epiphyte with thin tough leaves on pendulous stalks adapted to its rain forest habitat; or the rough-leaved sandpaper figs from Australia; or 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, often leading to very high levels of alpha diversity. In the tropics, it is quite common to find that Ficus is the most species-rich plant genus in a particular forest. In Asia as many as 70 or more species can co-exist.^[1]

Although identifying many of the species can be difficult, figs as a group are relatively easy to recognize. Often the presence of aerial roots or the general shape of the plant will give them away. Their fruit are also distinct. The fig fruit is in fact an enclosed inflorescence, sometimes referred to as a syconium, an urn-like structure lined on the inside with the fig's tiny flowers. The unique fig pollination system, involving tiny, highly specific wasps, know as fig wasps that enter these closed inflorescences to both pollinate and lay their own eggs, has been a constant source of inspiration and wonder to biologists^[2]. Finally, there are three vegetative traits that together are unique to figs. All figs possess a white to yellowish sap (latex), some in copious quantities; the twig has paired stipules or a circular stipule scar if the stipules have fallen off; and the lateral veins at the base of the leaf are steep, that is they form a tighter angle with the midrib than the other lateral veins, a feature referred to as a "tri-veined".

Unfortunately, there are no unambiguous older fossils of Ficus. However, current molecular clock estimates indicate that Ficus is a relatively ancient genus being at least 60 million years old^[2], 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.

Ecology and uses

Figs are keystone species in many rainforest ecosystems. Their fruit are a key resource for some frugivores including fruit bats, capuchin monkeys, langurs and mangabeys. They are even more important for some birds. Asian barbets, pigeons, hornbills, fig-parrots and bulbuls are examples of taxa which may almost entirely subsist on figs when these are in plenty. Many Lepidoptera caterpillars, for example of 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 feed on fig leaves. The Citrus long-horned beetle (Anoplophora chinensis), for example, has larvae which feed on wood, including that of fig trees; it 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 is 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).

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. Pou (F. religiosa leaves' shape inspired one of the standard kbach rachana, decorative elements in Cambodian architecture. Weeping Fig (F. benjamina) and Indian Rubber Plant (F. elastica) are identified as powerful air-cleaning plants in the NASA Clean Air Study. Indian Banyan (F. bengalensis) and the Indian Rubber Plant, as well as other species, have use in herbalism. The latter is known to be a hyperaccumulator of benzene and methane, and urban or potted plants should be considered toxic for that reason.

Figs have figured prominently in some human cultures. There is evidence that figs, specifically the Common Fig (F. carica) and Sycamore Fig (F. sycomorus), were among the first - if not the very 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 BC were found in the early Neolithic village Gilgal I (in the Jordan Valley, 13 km north of Jericho). These were a parthenocarpic type and thus apparently an early cultivar. This find predates the cultivation of grain in the Middle East many hundreds of years.^[3].

Cultural and spiritual significance

Fig trees has profoundly influenced culture through several religious traditions. Among the more famous species are the Sacred Fig tree (Peepul, Bodhi, Bo, or Po, Ficus religiosa) and the Banyan Fig (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 BC. It is one of the two sacred trees of Islam, and there is a sura in Quran named "The Fig" or At-Tin(سوره تین), and in East Asia, figs are pivotal in Buddhism, Hinduism and Jainism. Siddhārtha Gautama, the Supreme Buddha, is traditionally held to have found bodhi (enlightenment) while meditating under a Sacred Fig (F. religiosa). The same species was Ashvastha, 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 seems to be a Wavy-leaved Fig (F. infectoria). The Common Fig tree is cited in the Bible, where in Genesis 3:7, Adam and Eve cover their nakedness with fig leaves. The fig fruit is also included in the list of food found in the Promised Land, according to the Torah (Deut. 8). Other important plants reported included: wheat, barley, grapes, pomegranates, olives, and dates (representing the honey).

Fig pollination and fig fruit

See also: Common Fig

Many are grown for their fruits, though only Ficus carica is cultivated to any extent for this purpose. Furthermore, the fig fruits, important as both food and traditional medicine, contain laxative substances, flavonoids, sugars, vitamins A and C, acids and enzymes. However, figs are skin allergens, and the sap is a serious eye irritant. The fig is commonly thought of as fruit, but it is properly the flower of the fig tree. It is in fact a false fruit or multiple fruit, in which the flowers and seeds grow together to form a single mass. The genus Dorstenia, also in the figs family (Moraceae), exhibits similar tiny flowers arranged on a receptacle but in this case the receptacle is a more or less flat, open surface.

Figs, fresh Nutritional value per 100 g (3.5 oz)
Energy 70 kcal   310 kJ
Carbohydrates     19 g - Sugars  16 g - Dietary fiber  3 g   Fat 0.3 g Protein 0.8 g
Percentages are relative to US recommendations for adults. Source: USDA Nutrient database

Figs, dried Nutritional value per 100 g (3.5 oz)
Energy 250 kcal   1040 kJ
Carbohydrates     64 g - Sugars  48 g - Dietary fiber  10 g   Fat 1 g Protein 3 g
Percentages are relative to US recommendations for adults. Source: USDA Nutrient database

A fig "fruit" is derived from a specially adapted type of inflorescence (an arrangement of multiple flowers). In this case, it is an involuted, nearly closed receptacle with many small flowers arranged on the inner surface. Thus the actual flowers of the fig are unseen unless the fig is cut open. In Chinese the fig is called wú huā guǒ )Traditional Chinese: 無花果; Simplified Chinese: 无花果; ), "fruit without flower"^[4]. In Bengali, where the Common Fig is called dumur, it is referenced in a proverb: tumi jeno dumurer phool hoe gele ("You have become [invisible like] the dumur flower").

The syconium often 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 that crawl through the opening in search of a suitable place to lay eggs. Without this pollinator service fig trees cannot 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. Technically, a fig fruit proper would be one of the many tiny mature, seed-bearing flowers found inside one fig - if you cut open a fresh fig, the flowers will appear as fleshy "threads", each bearing a single seed inside.

The fig plants can be monoecious (hermaphrodite) or dioecious (hermaphrodite and female):^[5]

• 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).
• On the other hand the Common Fig (Ficus carica) is a dioecious plant, as well as 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 fig it grew up in. When the wasp dies, it is broken down by enzymes 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. Common Fig caprifigs have three crops per year; edible figs have two. The first (breba^[6]) produces small fruits called olynth. 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 or fig wasps.

There is typically only one species of wasp capable of fertilizing the flowers of each species of fig, 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 have been introduced, so only four species of figs produce viable seeds there. This is an example of mutualism, i.e. one organism (fig plant) can not propagate itself without the other one (fig wasp).

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^[7]. Additionally, recent genetic and molecular dating analyses have shown a very close correspondence in the character evolution and speciation phylogenies of these two clades.^[2]

Selected species

List of famous fig trees

• Ashvastha – the world tree of Hinduism, held to be a supernatural F. religiosa
• Bodhi tree – a F. religiosa
• Charybdis Fig Tree of the 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 probably F. infectoria
• Santa Barbara's Moreton Bay Fig Tree – a F. macrophylla
• Sri Maha Bodhi – another F. religiosa. Planted in 288 BC, the oldest human-planted tree on record
• 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

See also

Abraham Mauricio Salazar, famous papel amate artist Amphoe Pho Sai and Amphoe Suan Phueng, districts in Thailand named after Ficus species Banyan Edred John Henry Corner Fig Newton Fig-parrots

Figtree List of fruits Miracles of Jesus: the parable of the barren fig tree Mission fig Naturopathic medicine Nutrition

Pharmacosycea, a Ficus subgenus Phytonutrients Pippalada - Atharva-Veda scholar whose name means "Sacred Fig eater" Strangler Fig Figs in the Bible

Footnotes

References

• Berg, C.C. & Corner, E.J.H. (2005): Moraceae. In: Flora Malesiana Ser. I, vol. 17, part 2.
• California Rare Fruit Growers, Inc. (CRFG) (1996): Fig. Retrieved 2008-NOV-01.
• Carauta, Pedro; Diaz, Ernani (2002): Figueiras no Brasil. Editora UFRJ, Rio de Janeiro. ISBN 8571082502
• Denisowksi, Paul (2007): Chinese-English Dictionary – Fig. Retrieved 2008-NOV-01.
• Harrison, Rhett D. (2005): Figs and the diversity of tropical rain forests. Bioscience 55(12): 1053-1064. DOI:10.1641/0006-3568(2005)055[1053:FATDOT]2.0.CO;2 PDF fulltext
• Kislev, Mordechai E.; Hartmann, Anat & Bar-Yosef, Ofer (2006a): Early Domesticated Fig in the Jordan Valley. Science 312(5778): 1372. doi:10.1126/science.1125910 (HTML abstract) Supporting Online Material
• Kislev, Mordechai E.; Hartmann, Anat & Bar-Yosef, Ofer (2006b): Response to Comment on "Early Domesticated Fig in the Jordan Valley". Science 314(5806): 1683b. doi:10.1126/science.1133748 PDF fulltext
• Lev-Yadun, Simcha; Ne'eman, Gidi; Abbo, Shahal & Flaishman, Moshe A. (2006): Comment on "Early Domesticated Fig in the Jordan Valley". Science 314(5806): 1683a. doi:10.1126/science.1132636 PDF fulltext
• Lewington, Anna & Parker, Edward (1999): Ancient trees: Trees that live for 1000 years: 192. London, Collins & Brown Limited.
• Rønsted, Nina; Weiblen, George D.; Cook, James M.; Salamin, Nicholas; Machado, Carlos A. & Savoainen, Vincent (2005): 60 million years of co-divergence in the fig-wasp symbiosis. Proc. R. Soc. B 272(1581): 2593-2599. doi:10.1098/rspb.2005.3249 PDF fulltext
• Shanahan, M.; Compton, S.G.; So, Samson & Corlett, Richard (2001): Fig-eating by vertebrate frugivores: a global review. Biological Reviews 76(4): 529-572. doi:10.1017/S1464793101005760 PDF fulltext Electonic appendices

External links

• Figweb Major reference site for the genus Ficus
• Video: Interaction of figs and fig wasps Multi-award-winning documentary
• Fruits of Warm Climates: Fig
• Ficus Tree and Fig Care Information
• California Rare Fruit Growers: Fig Fruit Facts
• North American Fruit Explorers: Fig
• BBC: Fig fossil clue to early farming