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''Ficus aurea'', commonly known as the Florida strangler fig (or simply strangler fig), golden fig, or ''higuerón'', is a tree in the family
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 ...
that is native to the U.S. state of
Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
, the northern and western
Caribbean The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered around the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean, mostly overlapping with the West Indies. Bordered by North America to the north, Central America ...
, southern
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
and
Central America Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...
south to
Panama Panama, officially the Republic of Panama, is a country in Latin America at the southern end of Central America, bordering South America. It is bordered by Costa Rica to the west, Colombia to the southeast, the Caribbean Sea to the north, and ...
. The specific epithet ''aurea'' was applied by English botanist
Thomas Nuttall Thomas Nuttall (5 January 1786 – 10 September 1859) was an English botanist and zoologist who lived and worked in America from 1808 until 1841. Nuttall was born in the village of Long Preston, near Settle in the West Riding of Yorkshire a ...
who described the species in 1846. ''Ficus aurea'' is a strangler fig. In figs of this group, seed germination usually takes place in the canopy of a
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County * Host Island, in the Wilhelm Archipelago, Antarctica People * ...
tree with the seedling living as an
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 ...
until its roots establish contact with the ground. After that, it enlarges and strangles its host, eventually becoming a free-standing tree in its own right. Individuals may reach in height. Like all figs, it has an obligate mutualism with
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: figs are only pollinated by fig wasps, and fig wasps can only reproduce in fig flowers. The tree provides habitat, food and shelter for a host of tropical lifeforms including epiphytes in
cloud forests A cloud forest, also called a water forest, primas forest, or tropical montane cloud forest, is a generally tropical or subtropical, evergreen, montane, moist forest characterized by a persistent, frequent or seasonal low-level cloud cover, ...
and
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,
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s,
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s and
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s. ''F. aurea'' is used in
traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) refers to the knowledge, skills, and practices rooted in the cultural beliefs of various societies, especially Indigenous groups, used for maintaining health and treatin ...
, for live fencing, as an
ornamental Ornamental may refer to: *Ornamental grass, a type of grass grown as a decoration *Ornamental iron, mild steel that has been formed into decorative shapes, similar to wrought iron work *Ornamental plant, a plant that is grown for its ornamental qua ...
and as a
bonsai Bonsai (; , ) is the Japanese art of Horticulture, growing and shaping miniature trees in containers, with a long documented history of influences and native Japanese development over a thousand years, and with unique aesthetics, cultural hist ...
.


Description

''Ficus aurea'' is a tree which may reach heights of . It is
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 ...
: each tree bears functional male and female flowers. The size and shape of the leaves is variable. Some plants have leaves that are usually less than 10 cm (4 in) long while others have leaves that are larger. The shape of the leaves and of the leaf base also varies—some plants have leaves that are oblong or elliptic with a wedge-shaped to rounded base, while others have heart-shaped or ovate leaves with
cordate Cordate is an adjective meaning ' heart-shaped' and is most typically used for: * Cordate (leaf shape), in plants * Cordate axe, a prehistoric stone tool See also * Chordate A chordate ( ) is a bilaterian animal belonging to the phyl ...
to rounded bases. ''F. aurea'' has paired figs which are green when unripe, turning yellow as they ripen. They differ in size (0.6–0.8 cm .2–0.3 in about 1 cm .4 in or 1.0–1.2 cm .4–0.5 inin diameter); figs are generally sessile, but in parts of northern
Mesoamerica Mesoamerica is a historical region and cultural area that begins in the southern part of North America and extends to the Pacific coast of Central America, thus comprising the lands of central and southern Mexico, all of Belize, Guatemala, El S ...
figs are borne on short stalks known as peduncles.


Taxonomy

With about 750 species, ''
Ficus ''Ficus'' ( or ) is a genus of about 850 species of woody trees, shrubs, vines, epiphytes and hemiepiphytes in the family (biology), family Moraceae. Collectively known as fig trees or figs, they are native throughout the tropics with a few spe ...
'' (
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 ...
) is one of the largest
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit ...
genera (David Frodin of
Chelsea Physic Garden The Chelsea Physic Garden was established as the Apothecaries' Garden in London, England, in 1673 by the Worshipful Society of Apothecaries to grow plants to be used as medicines. This four acre physic garden, the term here referring to the scie ...
ranked it as the 31st largest genus). ''Ficus aurea'' is classified in the
subgenus In biology, a subgenus ( subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between the ge ...
''Urostigma'' (the strangler figs) and the
section Section, Sectioning, or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...
''Americana''. Recent molecular
phylogenies A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA. In o ...
have shown that subgenus ''Urostigma'' is
polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
, but have strongly supported the validity of section ''Americana'' as a
discrete group In mathematics, a topological group ''G'' is called a discrete group if there is no limit point in it (i.e., for each element in ''G'', there is a neighborhood which only contains that element). Equivalently, the group ''G'' is discrete if and ...
(although its exact relationship to section ''Galoglychia'' is unclear).
Thomas Nuttall Thomas Nuttall (5 January 1786 – 10 September 1859) was an English botanist and zoologist who lived and worked in America from 1808 until 1841. Nuttall was born in the village of Long Preston, near Settle in the West Riding of Yorkshire a ...
described the species in the second volume of his 1846 work ''The North American Sylva'' with specific epithet ''aurea'' ('golden' in
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
). In 1768, Scottish botanist
Philip Miller Philip Miller Royal Society, FRS (1691 – 18 December 1771) was an English botany, botanist and gardener of Scottish descent. Miller was chief gardener at the Chelsea Physic Garden for nearly 50 years from 1722, and wrote the highly popular ...
described ''
Ficus maxima ''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 176 ...
'', citing
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
' ''
Hortus Cliffortianus The ''Hortus Cliffortianus'' is a work of early botanical literature published in 1737. The work was a collaboration between Carl Linnaeus and the illustrator Georg Dionysius Ehret, financed by George Clifford in 1735–1736. Clifford was a we ...
'' (1738) and
Hans Sloane Sir Hans Sloane, 1st Baronet, (16 April 1660 – 11 January 1753), was an Irish physician, naturalist, and collector. He had a collection of 71,000 items which he bequeathed to the British nation, thus providing the foundation of the British ...
's ''Catalogus plantarum quæ in insula Jamaica'' (1696). Sloane's illustration of the species, published in 1725, depicted it with figs borne singly, a characteristic of the ''Ficus'' subgenus '' Pharmacosycea''. As a member of the subgenus '' Urostigma'', ''F. aurea'' has paired figs. However, a closer examination of Sloane's description led Cornelis Berg to conclude that the illustration depicted a member of the subgenus ''Urostigma'' (since it had other diagnostic of that subgenus), almost certainly ''F. aurea'', and that the illustration of singly borne figs was probably
artistic license Artistic license (and more general or contextually-specific, derivative terms such as creative license, poetic license, historical license, dramatic license, and narrative license) refers to deviation from fact or form for artistic purposes. It ...
. Berg located the plant collection upon which Sloane's illustration was based and concluded that Miller's ''F. maxima'' was, in fact, ''F. aurea''. In his description of ''F. aurea'', which was based on plant material collected in Florida, Thomas Nuttall considered the possibility that his plants belonged to the species that Sloane had described, but came to the conclusion that it was a new species. Under the rules of
botanical nomenclature Botanical nomenclature is the formal, scientific naming of plants. It is related to, but distinct from taxonomy (biology), taxonomy. Plant taxonomy is concerned with grouping and classifying plants; Botany, botanical nomenclature then provides na ...
, the name ''F. maxima'' has priority over ''F. aurea'' since Miller's description was published in 1768, while Nuttall's description was published in 1846. In their 1914 ''Flora of Jamaica'', William Fawcett and
Alfred Barton Rendle Alfred Barton Rendle FRS (19 January 1865 – 11 January 1938) was an English botanist. Rendle was born in Lewisham to John Samuel and Jane Wilson Rendle. He was educated in Lewisham where he first became interested in plants, St Olave's G ...
linked Sloane's illustration to the tree species that was then known as ''Ficus suffocans'', a name that had been assigned to it in
August Grisebach August Heinrich Rudolf Grisebach (; 17 April 18149 May 1879) was a German botanist and phytogeographer. Biography Grisebach studied at the Lyceum in Hanover, the cloister-school at Ilfeld, and the University of Göttingen. He graduated in me ...
's ''Flora of the British West Indian Islands''. Gordon DeWolf agreed with their conclusion and used the name ''F. maxima'' for that species in the 1960 ''Flora of Panama''.DeWolf, Gordon P., Jr. "Ficus (Tourn.) L." in Since this use has become widespread, Berg proposed that the name ''Ficus maxima'' be conserved in the way DeWolf had used it, a proposal that was accepted by the nomenclatural committee. Reassigning the name ''Ficus maxima'' did not leave ''F. aurea'' as the oldest name for this species, as German naturalist
Johann Heinrich Friedrich Link Johann Heinrich Friedrich Link (2 February 1767 – 1 January 1851) was a German natural history, naturalist and botanist. Biography Link was born at Hildesheim as a son of the minister August Heinrich Link (1738–1783), who taught him love ...
had described ''Ficus ciliolosa'' in 1822. Berg concluded that the species Link described was actually ''F. aurea'', and since Link's description predated Nuttall's by 24 years, priority should have been given to the name ''F. ciliolosa''. Since the former name was widely used and the name ''F. ciliolosa'' had not been, Berg proposed that the name ''F. aurea'' be conserved. In response to this, the nomenclatural committee ruled that rather than conserving ''F. aurea'', that it would be better to reject ''F. ciliolosa''. Conserving ''F. aurea'' would mean that precedence would be given to that name over all others. By simply rejecting ''F. ciliolosa'', the committee left open the possibility that the name ''F. aurea'' could be supplanted by another older name, if one were to be discovered.


Synonyms

In 1920, American botanist Paul C. Standley described three new species based on collections from Panama and Costa Rica—''Ficus tuerckheimii'', ''F. isophlebia'' and ''F. jimenezii''. DeWolf concluded that they were all the same species, and Berg synonymised them with ''F. aurea''. These names have been used widely for Mexican and Central American populations, and continue to be used by some authors. Berg suspected that ''Ficus rzedowskiana'' Carvajal and Cuevas-Figueroa may also belong to this species, but he had not examined the original material upon which this species was based. Berg considered ''F. aurea'' to be a species with at least four morphs. "None of the morphs", he wrote, "can be related to certain habitats or altitudes." Thirty years earlier, William Burger had come to a very different conclusion with respect to ''Ficus tuerckheimii'', ''F. isophlebia'' and ''F. jimenezii''—he rejected DeWolf's synonymisation of these three species as based on incomplete evidence. Burger noted that the three taxa occupied different habitats which could be separated in terms of rainfall and elevation.


Reproduction and growth

Figs have an obligate mutualism with
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, (Agaonidae); figs are only pollinated by these wasps, and they can only reproduce in fig flowers. Generally, each fig species depends on a single species of wasp for pollination. The wasps are similarly dependent on their fig species in order to reproduce. ''Ficus aurea'' is pollinated by '' Pegoscapus mexicanus'' ( Ashmead). Figs have complicated
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 ...
s called syconia. Flowers are entirely contained within an enclosed structure. Their only connection with the outside is through a small pore called
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 ...
.
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 like ''F. aurea'' have both male and female flowers within the syconium. Female flowers mature first. Once mature, they produce a volatile chemical attractant. Female wasps squeeze their way through the ostiole into the interior of the syconium. Inside the syconium, they pollinate the flowers, lay their eggs in some of them, and die. The eggs hatch and the larvae parasitise the flowers in which they were laid. After four to seven weeks (in ''F. aurea''), adult wasps emerge. Males emerge first, mate with the females, and cut exit holes through the walls of the fig. The male flowers mature around the same time as the female wasps emerge. The newly emerged female wasps actively pack their bodies with pollen from the male flowers before leaving through the exit holes the males have cut and fly off to find a syconium in which to lay their eggs. Over the next one to five days, figs ripen. The ripe figs are eaten by various mammals and birds which disperse the seeds.


Phenology

Figs flower and fruit asynchronously. Flowering and fruiting is staggered throughout the population. This fact is important for fig wasps—female wasps need to find a syconium in which to lay their eggs within a few days of emergence, something that would not be possible if all the trees in a population flowered and fruited at the same time. This also makes figs important food resources for
frugivore 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 ...
s (animals that feed nearly exclusively on fruit); figs are one of the few fruit available at times of the year when fruit are scarce. Although figs flower asynchronously as a population, in most species flowering is synchronised within an individual. Newly emerged female wasps must move away from their natal tree in order to find figs in which to lay their eggs. This is to the advantage of the fig, since it prevents
self-pollination Self-pollination is a form of pollination in which pollen arrives at the stigma of a flower (in flowering plants) or at the ovule (in gymnosperms) of the same plant. The term cross-pollination is used for the opposite case, where pollen from ...
. In Florida, individual ''F. aurea'' trees flower and fruit asynchronously. Within-tree asynchrony in flowering is likely to raise the probability of self-pollination, but it may be an adaptation that allows the species to maintain an adequate population of wasps at low population densities or in strongly seasonal climates. Flowering
phenology Phenology is the study of periodic events in biological life cycles and how these are influenced by seasonal and interannual variations in climate, as well as habitat factors (such as elevation). Examples include the date of emergence of leav ...
in ''Ficus'' has been characterised into five phases. In most figs, phase A is followed almost immediately by phase B. However, in ''F. aurea'' immature inflorescences can remain dormant for more than nine months.


Growth

''Ficus aurea'' is a fast-growing tree. As a hemiepiphyte it germinates in the
canopy Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape), aboveground portion of grapes Religion and ceremonies * Baldachin or canopy of state, typically placed over an a ...
of a host tree and begins life as an
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 ...
before growing roots down to the ground. ''F. aurea'' is also a strangler fig (not all hemiepiphytic figs are stranglers)—the roots fuse and encircle the host tree. This usually results in the death of the host, since it effectively girdles the tree. Palms, which lack
secondary growth In botany, secondary growth is the growth that results from cell division in the cambia or lateral meristems and that causes the stems and roots to thicken, while primary growth is growth that occurs as a result of cell division at the tips ...
, are not affected by this, but they can still be harmed by competition for light, water and nutrients. Following
Hurricane Andrew Hurricane Andrew was a compact, but very powerful and devastating tropical cyclone that struck the Bahamas, Florida, and Louisiana in August 1992. It was the most destructive hurricane to ever hit Florida in terms of structures dama ...
in 1992, ''F. aurea'' trees regenerated from root suckers and standing trees.


Distribution

''Ficus aurea'' ranges from Florida, across the northern Caribbean to Mexico, and south across Central America. It is present in central and southern Florida and the
Florida Keys The Florida Keys are a coral island, coral cay archipelago off the southern coast of Florida, forming the southernmost part of the continental United States. They begin at the southeastern coast of the Florida peninsula, about south of Miami a ...
, The Bahamas, the Caicos Islands, Hispaniola, Cuba, Jamaica, the Cayman Islands, San Andrés (a Colombian possession in the western Caribbean), southern Mexico, Belize, Guatemala, Honduras, Nicaragua, El Salvador, Costa Rica and Panama.Ficus aurea Nutt.
Flora Mesoamericana: Lista Anotada. Retrieved on 2008-07-02
It grows from sea level up to 1,800 m (5,500 ft)
above sea level Height above mean sea level is a measure of a location's vertical distance (height, elevation or altitude) in reference to a vertical datum based on a historic mean sea level. In geodesy, it is formalized as orthometric height. The zero level ...
, in habitats ranging from
Bahamian dry forests The Bahamian dry forests are a tropical and subtropical dry broadleaf forest ecoregion in the Bahamas and the Turks and Caicos Islands, covering an area of . They are found on much of the northern Bahamas, including Andros, Abaco, and Grand B ...
, to
cloud forest A cloud forest, also called a water forest, primas forest, or tropical montane cloud forest, is a generally tropical or subtropical, evergreen, Montane forest, montane, Tropical and subtropical moist broadleaf forests, moist forest characteri ...
in Costa Rica. ''Ficus aurea'' is found in central and southern Florida as far north as
Volusia County Volusia County (, ) is a county located in the east-central part of the U.S. state of Florida between the St. Johns River and the Atlantic Ocean. As of the 2020 census, the county was home to 553,543 people, an increase of 11.9% from the 2 ...
; it is one of only two native fig species in Florida. The species is present in a range of south Florida ecosystems, including coastal hardwood hammocks,
cabbage palm Cabbage palm is a common name for several species of palms or palm-like plants: *'' Cordyline fruticosa'', a tropical tree native to Asia and Polynesia *'' Corypha utan'', an East Asian fan palm (including Northern Australia) *''Euterpe oleracea'', ...
hammocks, tropical hardwood hammocks and shrublands, temperate hardwood hammocks and shrublands and along watercourses. In The Bahamas, ''F. aurea'' is found in tropical dry forests on
North Andros North Andros is one of the 31 districts of the Bahamas. It is also the largest district (in area) in the country. It has some of the largest settlements on Andros Island and many churches as well. The population (2022 Census) is 4,069. Churches ...
,
Great Exuma Exuma is a district of The Bahamas, consisting of over 365 islands and cays. The largest of the islands is Great Exuma, which is 37 mi (60 km) in length and joined to another island, Little Exuma, by a small bridge. The capital and largest to ...
and
Bimini Bimini is the westernmost district of the Bahamas and comprises a chain of islands located about due east of Miami. Bimini is the closest point in the Bahamas to the mainland United States and approximately west-northwest of Nassau. The popula ...
. ''F. aurea'' occurs in 10
states State most commonly refers to: * State (polity), a centralized political organization that regulates law and society within a territory **Sovereign state, a sovereign polity in international law, commonly referred to as a country **Nation state, a ...
in Mexico, primarily in the south, but extending as far north as
Jalisco Jalisco, officially the Free and Sovereign State of Jalisco, is one of the 31 states which, along with Mexico City, comprise the 32 Political divisions of Mexico, Federal Entities of Mexico. It is located in western Mexico and is bordered by s ...
. It is found in tropical deciduous forest, tropical semi-evergreen forest, tropical evergreen forest, cloud forest and in aquatic or subaquatic habitats.


Ecology

''Ficus aurea'' is a strangler fig—it tends to establish on a host tree which it gradually encircles and "strangles", eventually taking the place of that tree in the forest canopy. While this makes ''F. aurea'' an agent in the mortality of other trees, there is little to indicate that its choice of hosts is species specific. However, in
dry forests The tropical and subtropical dry broadleaf forest is a habitat type defined by the World Wide Fund for Nature and is located at tropical and subtropical latitudes. Though these forests occur in climates that are warm year-round, and may receive ...
on
Great Exuma Exuma is a district of The Bahamas, consisting of over 365 islands and cays. The largest of the islands is Great Exuma, which is 37 mi (60 km) in length and joined to another island, Little Exuma, by a small bridge. The capital and largest to ...
in The Bahamas, ''F. aurea'' establishes exclusively on palms, in spite of the presence of several other large trees that should provide suitable hosts. Eric Swagel and colleagues attributed this to the fact that
humus In classical soil science, humus is the dark organic matter in soil that is formed by the decomposition of plant and animal matter. It is a kind of soil organic matter. It is rich in nutrients and retains moisture in the soil. Humus is the Lati ...
accumulates on the leaf bases of these palms and provides a relatively moist
microclimate A microclimate (or micro-climate) is a local set of atmosphere of Earth, atmospheric conditions that differ from those in the surrounding areas, often slightly but sometimes substantially. The term may refer to areas as small as a few square m ...
in a dry environment, facilitating seedling survival. Figs are sometimes considered to be potential
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 communities of fruit-eating animals because of their
asynchronous Asynchrony is any dynamic far from synchronization. If and as parts of an asynchronous system become more synchronized, those parts or even the whole system can be said to be in sync. Asynchrony or asynchronous may refer to: Electronics and com ...
fruiting patterns. Nathaniel Wheelwright reports that emerald toucanets fed on unripe ''F. aurea'' fruit at times of fruit scarcity in
Monteverde Monteverde is the twelfth Cantons of Costa Rica, canton of the Puntarenas Province, Puntarenas Provinces of Costa Rica, province of Costa Rica, located in the Cordillera de Tilarán (Tilarán range). Roughly a four-hour drive from the Costa Rica ...
, Costa Rica. Wheelwright listed the species as a year-round food source for the
resplendent quetzal The resplendent quetzal (''Pharomachrus mocinno'') is a small bird found in Central America and southern Mexico that lives in tropical forests, particularly montane cloud forests. They are part of the family Trogonidae and have two recognized s ...
at the same site. In the
Florida Keys The Florida Keys are a coral island, coral cay archipelago off the southern coast of Florida, forming the southernmost part of the continental United States. They begin at the southeastern coast of the Florida peninsula, about south of Miami a ...
, ''F. aurea'' is one of five fruit species that dominate the diet fed by
white-crowned pigeon The white-crowned pigeon (''Patagioenas leucocephala'') is a fruit and seed-eating species of bird in the dove and pigeon family Columbidae. It is found primarily in the Caribbean. John James Audubon painted these pigeons, including the waterco ...
s to their nestlings. ''F. aurea'' is also important in the diet of mammalian frugivores—both fruit and young leaves are consumed by black howler monkeys in Belize. The interaction between figs 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, ...
s is especially well-known (see section on
reproduction Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. There are two forms of reproduction: Asexual reproduction, asexual and Sexual ...
, above). In addition to its pollinators (''Pegoscapus mexicanus''), ''F. aurea'' is exploited by a group of non-pollinating chalcidoid wasps whose larvae develop in its figs. These include
gall Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants. Plant galls are abnormal outgrowths of plant tissues, similar to benign tumors or war ...
ers,
inquiline In zoology, an inquiline (from Latin ''inquilinus'', "lodger" or "tenant") is an animal that lives commensally in the nest, burrow, or dwelling place of an animal of another species. For example, some organisms, such as insects, may live in the ...
s and
kleptoparasite Kleptoparasitism (originally spelt clepto-parasitism, meaning "parasitism by theft") is a form of feeding in which one animal deliberately takes food from another. The strategy is Evolutionarily stable strategy, evolutionarily stable when stealin ...
s as well as
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s of both the pollinating and non-pollinating wasps. The invertebrates within ''F. aurea'' syconia in southern Florida include a pollinating wasp, ''P. mexicanus'', up to eight or more species of non-pollinating wasps, a plant-parasitic nematode transported by the pollinator, mites, and a predatory rove beetle whose adults and larvae eat fig wasps. Nematodes: ''Schistonchus aureus'' (Aphelenchoididae) is a plant-parasitic nematode associated with the pollinator ''Pegoscapus mexicanus'' and syconia of ''F. aurea''. Mites: belonging to the family Tarsonemidae (Acarina) have been recognized in the syconia of ''F. aurea'' and ''F. citrifolia'', but they have not been identified even to genus, and their behavior is undescribed. Rove beetles: ''Charoxus spinifer'' is a rove beetle (Coleoptera: Staphylinidae) whose adults enter late-stage syconia of ''F. aurea'' and ''F. citrifolia''. Adults eat fig wasps; larvae develop within the syconia and prey on fig wasps, then pupate in the ground. As a large tree, ''F. aurea'' can be an important host for
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. In Costa Rican cloud forests, where ''F. aurea'' is "the most conspicuous component" of intact forest, trees in forest patches supported richer communities of epiphytic
bryophyte Bryophytes () are a group of embryophyte, land plants (embryophytes), sometimes treated as a taxonomic Division (taxonomy), division referred to as Bryophyta ''Sensu#Common qualifiers, sensu lato'', that contains three groups of non-vascular pla ...
s, while isolated trees supported greater
lichen A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...
cover.
Florida International University Florida International University (FIU) is a public research university with its main campus in Westchester, Florida, United States. Founded in 1965 by the Florida Legislature, the school opened to students in 1972. FIU is the third-largest univ ...
ecologist Suzanne Koptur reported the presence of
extrafloral nectaries Nectar is a viscous, sugar-rich liquid produced by plants in glands called nectaries, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, ...
on ''F. aurea'' figs in the Florida
Everglades The Everglades is a natural region of flooded grasslands in the southern portion of the U.S. state of Florida, comprising the southern half of a large drainage basin within the Neotropical realm. The system begins near Orlando with the K ...
. Extrafloral nectaries are structures which produce
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
but are not associated with flowers. They are usually interpreted as defensive structure and are often produced in response to attack by insect
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
s. They attract insects, primarily
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s, which defend the nectaries, thus protecting the plant against herbivores.


Uses

''Ficus aurea'', amongst other related ''Ficus'' species, has been a source of bark for preparing ''
amate Amate ( from ) is a type of bark paper that has been manufactured in Mexico since the precontact times. It was used primarily to create codices. Amate paper was extensively produced and used for both communication, records, and ritual during t ...
'', the bark paper used for
codices The codex (: codices ) was the historical ancestor format of the modern book. Technically, the vast majority of modern books use the codex format of a stack of pages bound at one edge, along the side of the text. But the term ''codex'' is now r ...
in the Mesoamerican civilizations. The oldest example dates back to 75 CE and was found in a shaft tomb culture site in Huitzilapa,
Jalisco Jalisco, officially the Free and Sovereign State of Jalisco, is one of the 31 states which, along with Mexico City, comprise the 32 Political divisions of Mexico, Federal Entities of Mexico. It is located in western Mexico and is bordered by s ...
in
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
. The fruit of ''Ficus aurea'' is edible and was used for food by the Native Americans and early settlers in Florida; it is still eaten occasionally as a backyard source of native fruit. The latex was used to make a chewing gum, and aerial roots may have been used to make lashings, arrows, bowstrings and fishing lines. The fruit was used to make a rose-coloured dye. ''F. aurea'' was also used in
traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) refers to the knowledge, skills, and practices rooted in the cultural beliefs of various societies, especially Indigenous groups, used for maintaining health and treatin ...
in The Bahamas and Florida. Allison Adonizio and colleagues screened ''F. aurea'' for anti-
quorum sensing In biology, quorum sensing or quorum signaling (QS) is the process of cell-to-cell communication that allows bacteria to detect and respond to cell population density by gene regulation, typically as a means of acclimating to environmental disadv ...
activity (as a possible means of anti-bacterial action), but found no such activity. Individual ''F. aurea'' trees are common on dairy farms in La Cruz, Cañitas and Santa Elena in Costa Rica, since they are often spared when forest is converted to
pasture Pasture (from the Latin ''pastus'', past participle of ''pascere'', "to feed") is land used for grazing. Types of pasture Pasture lands in the narrow sense are enclosed tracts of farmland, grazed by domesticated livestock, such as horses, c ...
. In interviews, farmers identified the species as useful for fence posts, live fencing and firewood, and as a food species for wild birds and mammals. ''Ficus aurea'' is used as an
ornamental tree Ornamental plants or ''garden plants'' are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars th ...
, an indoor tree and as a
bonsai Bonsai (; , ) is the Japanese art of Horticulture, growing and shaping miniature trees in containers, with a long documented history of influences and native Japanese development over a thousand years, and with unique aesthetics, cultural hist ...
. Like other figs, it tends to invade built structures and foundations, and needs to be removed to prevent structural damage. Although young trees are described as "rather ornamental", older trees are considered to be difficult to maintain (because of the
adventitious roots Important structures in plant development are buds, shoots, roots, leaves, and flowers; plants produce these tissues and structures throughout their life from meristems located at the tips of organs, or between mature tissues. Thus, a living plant ...
that develop off branches) and are not recommended for small areas. However, it was considered a useful tree for "enviroscaping" to conserve energy in south Florida, since it is "not as aggressive as many exotic fig species," although it must be given enough space.


References


External links


Interactive Distribution Map for Ficus aurea
{{Taxonbar, from=Q2561277 aurea Trees of the Caribbean Trees of Central America Flora of Florida Trees of Northern America Plants described in 1846 Taxa named by Thomas Nuttall Epiphytes Garden plants of North America Plants used in bonsai Ornamental trees Flora without expected TNC conservation status