| Ficus insipida | |
|---|---|
 | |
| Roots | |
Scientific classification  | |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Angiosperms |
| Clade: | Eudicots |
| Clade: | Rosids |
| Order: | Rosales |
| Family: | Moraceae |
| Genus: | Ficus |
| Subgenus: | F. subg. Pharmacosycea |
| Species: | F. insipida |
| Binomial name | |
| Ficus insipida | |
| Synonyms [2] | |
Ficus insipida is a common tropical tree in the fig genus of the family Moraceae growing in forest habitats along rivers. It ranges from Mexico to northern South America. [3]
The tree was described in 1806 under the scientific name Ficus insipida (literally "insipid fig") by Carl Ludwig Willdenow, having studied the herbarium specimens collected in Caracas by the gardener Franz Bredemeyer in the 1780s during the Märter Expedition . Willdenow reports its fruit are tasteless. [4] Incongruously, among the many species of figs to grow in the region, this species is in fact recognisable by its large and sweet figs (when ripe). [5]
In the 1960 Flora of Panama, Gordon P. DeWolf Jr. lumped the species F. adhatodifolia and F. crassiuscula as synonyms of F. insipida, [6] but his taxonomic interpretation was not followed by subsequent authorities. [2]
Ficus expert Cees Berg distinguished two allopatric or almost allopatric subspecies in 1984: [7]
With about 750 species, Ficus (Moraceae) is one of the largest angiosperm genera. [9] F. insipida is classified in subgenus Pharmacosycea , section Pharmacosycea, subsection Bergianae (for which it is the type species), along with F. adhatodifolia, F. carchiana, F. crassiuscula, F. gigantosyce, F. lapathifolia, F. mutisii, F. oapana (spec. nov.? ined.), F. obtusiuscula, F. piresiana, F. rieberiana and F. yoponensis. [10] Although recent work suggests that subgenus Pharmacosycea is polyphyletic, section Pharmacosycea appears to be monophyletic and is a sister group to the rest of the genus Ficus. [11]
Mysteriously, genetic testing of a single individual of the three fig species F. maxima, F. tonduzii and F. yoponensis, each collected on Barro Colorado Island in Panama, found each species to be phylogenetically nestled within F. insipida. All four species occur together in a similar large range, but nonetheless, these species are quite easily differentiated morphologically. Strangely, in the case of the F. maxima and F. tonduzii specimens, these were nestled within a different haplotype, with an Amazonian distribution, as opposed to clustering within the haplotype found contemporaneously in Panama. An explanation for this is not readily apparent: the species may have recently evolved from F. insipida, although this would seem unlikely, or perhaps all three specimens just happened to be hybrids, another unlikely possibility. [12]
This is a tree with buttress roots that ranges from 8–40 m (26–131 ft) tall. [6] Because this is a pioneer species which quickly colonises secondary forest, and it is also a fast-growing species which can grow into a massive tree in only 100 years or so, it is generally readily recognisable as the largest trees in such secondary woodlands. [13]
Leaves vary shape from narrow to ellipse-shaped; they range from 5–25 cm (2.0–9.8 in) long and from 2–11 cm (0.79–4.33 in) wide. [6]
In Costa Rica or Panama it may be confused with Ficus yoponensis, but this similar lowland fig tree has smaller leaves, stipules and fruit, and only occurs in primary forest, whereas F. insipida is also found in secondary forest. [13] [14] [15] Another similar fig species in this area is F. crassivenosa, but this species has differently shaped leaves and does not have the same habitat preference (the tendency to grow in association with water courses). [8]
The nominate subspecies occurs from Mexico south throughout Central America to Colombia and Venezuela and thence to Ecuador, Bolivia, Peru and Amazonian Brazil, and north from Venezuela to Trinidad and Tobago and the Lesser Antilles, [8] [16] [17] whereas subspecies scabra occurs from the Guianan Shield of northeast Venezuela eastwards through the Guianas to northwestern Brazil in the states of Amapá and Pará. [7] [8]
In Mexico it has been recorded to occur in the states of Chihuahua, Durango, San Luis Potosí, Sinaloa, Sonora, Tamaulipas and Zacatecas in the north, southwards to Campeche, Chiapas, Colima, Guerrero, Hidalgo, Jalisco, México, Michoacán de Ocampo, Morelos, Nayarit, Oaxaca, Puebla, Querétaro, Tabasco and Veracruz de Ignacio de la Llave. [16] [17] In both Costa Rica and Nicaragua it is found in lowlands along both the Atlantic and Pacific coasts, as well as in the central valleys. [3] [8] [18]
In Bolivia it has been recorded in the northern and eastern departments of Beni, Cochabamba, La Paz, Pando and Santa Cruz: most of the country except the Andes in the southwest. In Ecuador it is known from the provinces of Esmeraldas, Imbabura, Manabí, Morona-Santiago, Napo, Pastaza, Sucumbíos and Zamora-Chinchipe. [17] In Colombia the species has been recorded in the departments of Amazonas, Antioquia, Bolívar, Boyacá, Caquetá, Casanare, Cauca, Chocó, Cundinamarca, La Guajira, Guaviare, Huila, Magdalena, Meta, Nariño, Norte de Santander, Putumayo, Risaralda, Santander, Tolima and Valle. [19]
The distribution in Brazil includes, besides Amapá and Pará mentioned above, the states of Acre, Amazonas and Rondônia. In the state of Pará both subspecies appear to occur, although this is unclear. [20]
Although it is often stated that the Amazon rainforest is ancient, much of it has in fact grown quite recently, after the end of the last Ice Age and with a large expansion to the south 3,000 years ago. During the Ice Age large tracts of the Amazon were covered in savanna, with the forest having retreated to numerous refugia. Traces of this were found in the genetic structure of the populations of the nominate subspecies: although populations are reasonably diverse from Mexico to the Andes area, the trees in the populations in most of the Amazon area are genetically similar to one another, with the occurrence of a "single widespread haplotype" and the trees in much of Bolivia having no discernable genetic diversity found in the tested sequences at all, indicating they only recently colonised this region. In this the species shows a similar pattern to other lowland, rainforest trees. [12]
The nominate subspecies is quite common in Nicaragua and Panama, [3] [13] but subspecies scabra is conversely said to be a rare tree in the Guianas. [5]
The typical habitat of the nominate subspecies is lowland forests down to the coast. [3] [8] Ficus insipida subsp. insipida does not grow above 1,100m in Costa Rica, [8] [18] and is found down to 50m, [18] or sea level. [8] It grows between 0–700m in altitude in Nicaragua (exceptionally up to 1300m). [3] It can be found in either very humid, humid or dry climates, but it is almost always encountered growing along rivers, and often on slopes. [3] [8] [18] It is found in thickly wooded small hills abutting the coast in Atlantic Costa Rica. [21]
The scabra subspecies appears to have a slightly different habitat preference, being typically found on slopes in either rainforest or mountain savannas in the Guianas. [5]
Like many figs and other rainforest fruit trees, F. insipida is a 'mass-fruiter', and like many (tropical rainforest) figs but unlike many rainforest fruit trees, individual F. insipida trees fruit according to a staggered, asynchronous schedule with respect to other neighbouring conspecifics. Thus, irrespective of the season, there is always a fig flowering and fruiting somewhere, which can be advantageous to wildlife, and functions to force its wasp pollinators to seek out a new tree, and thus foster cross pollination. Like the fruit, the new leaves are produced in asynchronous flushes. [22]
It is a monoecious species, [12] the figs, actually a specialised inflorescence called a synconium, are densely coated in minute flowers ('florets') on the inside, both functionally male and female. The female flowers mature first; they are found in two different versions, with a short or long style. The stigmas of the female flowers are thickly intertwined and coherent to each other at the same height (short-styled florets are simply positioned somewhat higher using pedicels and somewhat longer ovaries to maintain the stigma surface), and form a surface layer a certain distance from the inner wall of the fig, called the synstigma - this synstigma essentially functions as a platform on which the pollinating wasps must walk and from where they must oviposit their eggs. The synstigma is so coherent, pollen tubes may grow from one stigma into the ovule of another neighbouring floret. The length specified by the distance between the synstigma and the ovules helps determine which wasp species may live in a particular fig species, and also cause the females to mostly lay their eggs in the short-styled florets (although in F. insipida this is not so strict, and both types of florets are fertile and both can host a wasp larva). [10] [23]
The flowers found within the figs of F. insipida are pollinated by the females of tiny wasps belonging to the genus Tetrapus , which complete much of their lifecycle within the developing figs. The female wasps are weak-jawed, and rely on the males to free them from their figs and individual fruit in which they develop and pupate, but only the females are winged, and can thus fly to the next fig to lay their eggs. The males develop first, they are wingless but have stronger jaws, which they use to chew their way to freedom from their host ovule. Once free, they chew free the females, copulating with them while the females are still largely trapped in their ovules -this ensures each female has sex, females which do not copulate will only produce males as offspring. The males also chew holes through the walls of the fig and open up the ostiole (a small opening at the apex of the fig), allowing the females to escape. Meanwhile, the male flowers within the fig finally shed their pollen, which adhere to the females in specialised pockets or simply onto their body surface. The females search for a new fig in which to lay their eggs, and upon arriving upon one must embark upon their greatest challenge: forcing their way within through the ostiole. [12] [23] Although the wasps are quite minuscule, they nonetheless regularly undertake reasonable journeys, as can be seen in the genetic structure of the fig tree populations: there is clear evidence of abundant outcrossing in the nuclear DNA (which is transported in the pollen dusted on the females, as opposed to mitochondrial DNA). [12] The ostiole is barred by a series of bracts, but unlike in many other Ficus species, only the uppermost ostiolar bracts are interlocking and patent, with the inner bracts positioned inward and relatively open, thus forming a long slit-like tunnel allowing access to the central cavity. [10] [23] Nevertheless, entering the cavity is a strenuous task, and the females are often die in the tunnel, or are damaged by the ordeal, with their wings invariably torn off from forcing their way through the bracts. Once inside, the females inject their eggs with their ovipositor, through the styles of the correct length, into the ovules: one egg an ovule. While doing so, the females pollinate the other flowers when walking around on the synstigmatic surface. The seeds and the larvae mature in a few weeks, at approximately the same rate. [23]
It uses zoochory to disperse its seeds. [5] The figs are eaten by bats, howler, spider and capuchin monkeys in the Guianas. [5] An especially important species to aid in dispersal via endozoochory in Costa Rica is possibly the large and common trout-like fish Brycon guatemalensis , of which the adults primarily feed upon the fallen leaves and figs of F. insipida. The seeds can survive the passage through the gut of the fish, although their viability is significantly diminished. Nonetheless, the fish may have a specific value for the fig as a dispersal agent: this fig species primarily and typically is found along rivers, and fish have the advantage of generally dispersing the seeds along rivers. Furthermore, fish are able to disperse upriver, and thus maintain upriver populations, whereas dispersal by floating the figs in water (hydrochory) alone is generally in a downriver direction (in most habitats). [24] The leaves and especially the fruit of F. insipida and F. yoponensis are a preferred food of howler monkeys in Panama ( Alouatta palliata ), with one troop on Barro Colorado Island spending one quarter of its time feeding on these two types of trees. The asynchronous plant characteristics, and the food preference of the monkeys, results in the monkeys adopting specific foraging routes in order to check up on the status of as much of the potential trees as can be done efficiently. [22]
The latex is sold in South America as an anthelmintic, marketed as 'doctor oje' (ojé in Brazil). An analysis of overdoses in one area of Peru found only 3 cases that were apparently fatal and 39 cases requiring hospitalization over a 12 year period, which, based on an analysis of probable sales in the region, amounted to very low 0.01–0.015% fatality and 0.13–0.2% hospitalization rates. [25] The authors concluded the product was safe when dosed properly. No serious adverse effects were observed in any of several clinical trials on the product conducted in Peru, except for possibly one miscarriage in one 18-year-old woman (who did not disclose her pregnancy to those running the clinical trial) receiving a very low dose of ojé. [26]
The latex can be purified, leaving a complex of enzymes known as ficin, a white powder that was first produced in 1930. [27] This product is likely safe. [27] It was initially observed that intestinal nematodes dissolved in a ficin solution, which increased interest in the product at the time as an anthelmintic, although it was not widely adopted.
Ficin is a mix of different enzymes and can be produced from many different species of Ficus. The main proteolytic enzyme found in ficin produced from F. insipida has officially been named ficain. [28] Purified ficin has numerous medical and industrial uses. It is used for cleaning in the production of stitching material for sutures, to prepare animal arteries before transplantation into humans, [27] and for unmasking antigens in serology. [28] It is similarly used for cleaning the animal intestines used as sausage or cheese-casings. It is used as an additive to make freeze-resistant beer, and has been added to certain formulations of meat tenderizers along with related protease-type enzymes. [27]
According to Schultes and Raffauf in their 1990 book The Healing Forest, the fruit of Ficus anthelmintica (an antiquated synonym of F. insipida) has been used by an unknown people somewhere in the northern Amazon of Brazil as an aphrodisiac and for what they categorise as a 'memory enhancer'. [29]
As of 2021, the conservation status has not been assessed by the Centro Nacional de Conservação da Flora, [30] nor in the IUCN Red List, [31] nor by the Costa Rican national authority. In Costa Rica this species is present in numerous protected areas, such as Estación Biológica La Selva, Carara National Park, Corcovado National Park, [18] and Jairo Mora Sandoval Gandoca-Manzanillo Mixed Wildlife Refuge. [21]
Ficus 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, 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.
Ficus macrophylla, commonly known as the Moreton Bay fig or Australian banyan, is a large evergreen banyan tree of the Mulberry Family (Moraceae) native to eastern Australia, from the Wide Bay–Burnett region in the north to the Illawarra in New South Wales, as well as Lord Howe Island where the subspecies F. m. columnaris is a banyan form covering 2.5 acres or more of ground. Its common name is derived from Moreton Bay in Queensland, Australia. It is best known for its imposing buttress roots.
Ficus rubiginosa, the rusty fig or Port Jackson fig, is a species of flowering plant native to eastern Australia in the genus Ficus. Beginning as a seedling that grows on other plants (hemiepiphyte) or rocks (lithophyte), F. rubiginosa matures into a tree 30 m (100 ft) high and nearly as wide with a yellow-brown buttressed trunk. The leaves are oval and glossy green and measure from 4 to 19.3 cm long and 1.25 to 13.2 cm wide.
Ficus microcarpa, also known as Chinese banyan, Malayan banyan, Indian laurel, curtain fig, or gajumaru (ガジュマル), is a tree in the fig family Moraceae. It is native in a range from China through tropical Asia and the Caroline Islands to Australia. It is widely planted as a shade tree and frequently misidentified as F. retusa or as F. nitida.
Ficus citrifolia, also known as the shortleaf fig, giant bearded fig, Jagüey, wild banyantree and Wimba tree, is a species of banyan native to southern Florida, the Caribbean, Mexico, Central America, and northern South America south to Paraguay. It is distinguished from the closely related Florida strangler fig mainly by the finer veining in the leaves.
Syconium is the type of fruit borne by figs, 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 considered both a multiple and accessory fruit.
Ficain also known as ficin, debricin, or higueroxyl delabarre is a proteolytic enzyme extracted from the latex sap from the stems, leaves, and unripe fruit of the American wild fig tree Ficus insipida.
Ficus pumila var. awkeotsang, also known as the jelly fig, aiyu, or ai-yu, is a variety of Ficus pumila, and a member of the fig family Moraceae, native to Taiwan. The plant is known for its use in making aiyu jelly.
Ficus crassiuscula is a species of flowering plant in the family Moraceae, native to Central America and north-western parts of South America.
Ficus aurea, commonly known as the Florida strangler fig, golden fig, or higuerón, is a tree in the family Moraceae that is native to the U.S. state of Florida, the northern and western Caribbean, southern Mexico and Central America south to Panama. The specific epithet aurea was applied by English botanist Thomas Nuttall who described the species in 1846.
Ficus maxima is a fig tree which is native to Mexico, Central America, the Caribbean and South America south to Paraguay. Figs belong to the family Moraceae. The specific epithet maxima was coined by Scottish botanist Philip Miller in 1768; Miller's name was applied to this species in the Flora of Jamaica, but it was later determined that Miller's description was actually of the species now known as Ficus aurea. To avoid confusion, Cornelis Berg proposed that the name should be conserved for this species. Berg's proposal was accepted in 2005.
Pharmacosycea is one of six subgenera currently recognised in the genus Ficus. It was proposed by E. J. H. Corner in 1967 to unite section Pharmacosycea with Oreosycea.
Ficus americana, commonly known as the West Indian laurel fig or Jamaican cherry fig, is a tree in the family Moraceae which is native to the Caribbean, Mexico in the north, through Central and South America south to southern Brazil. It is an introduced species in Florida, USA. The species is variable; the five recognised subspecies were previously placed in a large number of other species.
Ficus obliqua, commonly known as the small-leaved fig, is a tree in the family Moraceae, native to eastern Australia, New Guinea, eastern Indonesia to Sulawesi and islands in the southwestern Pacific Ocean. Previously known for many years as Ficus eugenioides, it is a banyan of the genus Ficus, which contains around 750 species worldwide in warm climates, including the edible fig. Beginning life as a seedling, which grows on other plants (epiphyte) or on rocks (lithophyte), F. obliqua can grow to 60 m (200 ft) high and nearly as wide with a pale grey buttressed trunk, and glossy green leaves.
Ficus yoponensis is a species of fig tree found in Central and South America. It can grow to heights of 40–50 metres (130–160 ft) tall, having a trunk diameter of 1 metre (3.3 ft). The trunk is buttressed, light grey in colour and reasonably smooth. Its petioles are 1–2.5 centimetres (0.39–0.98 in) long, the stipules are straight and 3–5 centimetres (1.2–2.0 in) long. The leaves and stems are hairless. The leaves are 6–11 centimetres (2.4–4.3 in) long and 2.5–4 centimetres (0.98–1.57 in) wide, but larger in juveniles, being up to 28 centimetres (11 in) long and 5 centimetres (2.0 in) wide. The time at which they flower varies between individuals, but each tree tends to flower at a similar time each year. As in all figs, the flowers are enclosed inside the fig and can only be accessed by fig wasps, which enter to pollinate the flowers and lay their own eggs. The resulting fruit grows to 1.8 centimetres (0.71 in) in diameter and turns from green to purple with maturity. On average in Panama, F. yoponensis produce a new flush of leaves every 20 weeks and flower every 25 weeks. The species is similar in appearance to Ficus insipida but has smaller leaves, stipules and fruits and only occurs in primary forest whereas F. insipida is also found in secondary forest.
Ficus septica is a shrub or tree of the family Moraceae living at low altitudes from northeast India to north Australia (Queensland), and throughout Malesia. It lives on the edge of the vegetation, often in degraded environments. The seeds of this species are dispersed by numerous species, including fruit bats (Megachiroptera) when present.
Ficus hispida also known as the opposite leaf Fig is a small but well distributed species of tropical fig tree. It is morphologically gynodioecious, but functionally dioecious. Male trees are hermaphrodites with both staminate flowers that produce pollen and pistillate flowers that produce almost no seeds but can form galls containing pollinator wasp larvae. Female trees have pistillate flowers that do produce seeds but are inhospitable to pollinator wasp larvae.
Ficus variegata is a well distributed species of tropical fig tree. It occurs in many parts of Asia, islands of the Pacific and as far south east as Australia. There is a large variety of local common names including common red stem fig, green fruited fig and variegated fig. A non strangling fig which may reach 30 metres in height. The tree is evergreen when young but becomes briefly deciduous as it grows older. In Australia the fruit are eaten by cassowaries and double-eyed fig parrots.
Ficus burtt-davyi is a fig species endemic to Southern Africa, belonging to the Mulberry family of Moraceae. It grows in coastal and inland forests up to 1500m, from the vicinity of Mossel Bay in the Southern Cape to southern Mozambique - the forms growing on coastal dunes in the northern part of its range are salt tolerant and form low thickets on the margins of woodland. In the southern and eastern Cape forests the species becomes a strangler or liane, while when found on rocky outcrops and cliffs it usually develops into a rock-splitter.
Ficus bernaysii is a lowland rainforest tree in the family Moraceae, native to an area from New Guinea to the Solomon Islands. It is dioecious, and grows cauliflorous fruit. It is fed on by a wide range of animals.
