Ficus aurea
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| Florida strangler fig | ||||||||||||||
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| Ficus aurea Nutt. |
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Ficus ciliolosa Link |
Ficus aurea, commonly known as the Florida strangler fig (or simply strangler fig) or higuerón[1], is a species or species complex native to Florida, the northern Caribbean, southern Mexico, throughout Central America and on the Colombian island of San Andrés.[2] Originally described by Thomas Nuttall, its specific epithet is the Latin adjective aurea 'golden'.[3]
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[edit] Description
Ficus aurea is a monoecious[4] tree tree which grows up to 30 m (100 ft) tall.[5] Figs are generally evergreen, but at the northern end of its distribution in Florida, F. aurea is briefly leafless in winter.[4] 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 usually larger. The shape of the leaves and of the leaf base also varies - some plants have leaves that are oblong or elliptic with a cuneate to rounded base, while others have cordiform or ovate leaves with cordate to rounded bases. F. aurea has paired figs. They differ in size (0.6-0.8 cm, about 1 cm, or 1-1.2 cm in diameter) and can be either sessile or pedunculate.[2]
[edit] Reproduction and growth
Figs have an obligate mutualism with fig wasp (Agaonidae); figs are only pollinated by fig wasps, and fig wasps 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).[6]
Fig have complicated inflorescences called syconia. Flowers are entirely contained within an enclosed structure. Their only connection with the outside is through a small pore called ostiole. Monoecious figs have both male and female flowers within the syconium.[7] Female flowers mature first. Once mature they produce a volatile chemical attractant.[8] Female wasps squeeze their way through the ostiole into the interior of the syconium. Inside the syconium, the female wasp pollinates the flowers and lays her eggs and then dies. The eggs parasitise the flowers in which they are laid. After four to seven weeks (in F. aurea), the wasps emerge and the male flowers mature. Wasps mate within the fig. The males cut exit holes while the females pack pollen onto their bodies. Females leave 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.[8]
[edit] Phenology
Figs flower and fruit asynchronously.[4] Flowering and fruiting is staggered throughout the population. This fact is important for fig wasps - female wasps need to find a syconium to lay their eggs in within a few days of emergence, something that would not be possible if all the trees in a population flowered at fruited at the same time. This also makes figs important food resources for frugivores; 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.[7] In Florida, individual F. aurea trees flower and fruit asynchronously.[4] 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.[4]
- Flowering phenology in F. aurea
| Phase[8] | Description[8] | Duration in F. aurea[8] |
|---|---|---|
| A (pre-female) | Immature flowers | 2 days to >9 months |
| B (female) | Female flowers that are receptive to pollination; female wasps lay eggs and pollinate flowers | 1 day to 3 weeks |
| C (interfloral) | Fig seeds and wasp larvae develop | 4 to 7 weeks |
| D (male) | Male flowers mature; wasps emerge, mate and female wasps disperse | 1 to 2 days |
| E (post-floral) | Fruits ripen | 1 to 5 days |
Flowering phenology 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.[4]
[edit] Growth
Ficus aurea is a hemiepiphytic fig. It germinates in the canopy of a host tree and begin life as an epiphyte 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 tree (since it effectively girdles the tree). Palms, which lack secondary growth, are not affected by this, but they can still be harmed by competition for light, water and nutrients.[9] In Great Exuma, palms are the only hosts for stranglers, despite the presence of other large trees.[10]
Following Hurricane Andrew in 1992 F. aurea trees regenerated from root suckers and standing trees.[11]
[edit] 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[12], The Bahamas, the Caicos Islands, Hispaniola, Cuba, Jamaica, the Cayman Islands, San Andrés (a Colombian possession in the western Caribbean),[2] southern Mexico,[13] Belize, Guatemala, Honduras, Nicaragua, El Salvador, Costa Rica and Panama.[14]
Ficus aurea is found in central and southern Florida as far north as Volusia County.[15] The species is present in a number of south Florida ecosystems, including coastal hardwood hammocks, cabbage palm hammocks, tropical hardwood hammocks and shrublands, temperate hardwood hammocks and shrublands[16] and along watercourses.[8]
In The Bahamas, F. aurea is found in dry forests in North Andros[17], Great Exuma[10] and Bimini.[18]
In Mexico, F. aurea occurs in southern Mexico, as far north as Jalisco.[13] It is found in tropical deciduous forest, tropical semi-evergreen forest, tropical evergreen forest, cloud forest and in aquatic or subaquatic habitats.[13]
[edit] Ecology
Ficus aurea grows from sea level up to 1800 m (5500 ft) above sea level,[2] in habitats ranging from tropical dry forest in The Bahamas,[10] to cloud forest in Costa Rica.[19]
Figs are sometimes considered keystone species in frugivore communities because of their asynchronous fruiting patterns. Wheelwright reports that Emerald Toucanets fed on unripe F. aurea fruit at times of fruit scarcity in Monteverde, Costa Rica.[20] Wheelwright also listed it as a year-round food source for the Resplendent Quetzal at Monteverde.[21]
Suzanne Koptur reported the presence of extrafloral nectaries on F. aurea figs in the Florida Everglades.[22]
[edit] Uses
The fruit of Ficus aurea is edible and was used for food by the indigenous people and early settlers in Florida.[23] The latex was used to make a chewing gum, and aerial roots may have been used to make lashings, arrows, bowstrings and fishing lines.[23] The fruit was used to make a rose-coloured dye.[23] F. aurea was also used in traditional medicine in The Bahamas[24] and Florida.[23]
Allison Adonizio and colleagues screened F. aurea for anti-quorum sensing activity (as a possible means of anti-bacterial action), but found no such activity.[25]
Ficus aurea was a common remnant trees in diary farms in La Cruz, Cañitas and Santa Elena in Costa Rica. Farmers identified it as useful for fence posts, live fences and firewood, and as a food species for wild birds and mammals.[1]
[edit] Taxonomy
With about 750 species, Ficus (Moraceae) is one of the largest angiosperm genera. (Frodin ranked it in 31st place.)[26] Ficus aurea classified in the subgenus Urostigma (the strangler figs) and the section Americana.[2] Recent work has shown that subgenus Urostigma is polyphyletic, but that section Americana is strongly supported.[27]
This species was first described by Philip Miller as Ficus maxima in 1768, citing Linneaus' Hortus Cliffortianus (1738) and Hans Sloane's Catalogus plantarum quæ in insula Jamaica (1696). Sloane's illustration of this plant[1] (published in his 1725 A voyage to the islands Madera, Barbados, Nieves, S. Christophers and Jamaica) 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. Nuttall considered the possibility that his material from Florida belonged to the species that Sloane had illustrated, but decided that it must be a new species. However, a closer examination of Sloane's description led Cornelis Berg to conclude that the illustration was of 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". Berg also located the plant collection upon which Sloane's illustration was based, and concluded that it was F. aurea.[28]
When Thomas Nuttall described F. aurea based on collections from Florida, he considered the possibility that his plants belonged to the same species that Sloane had described, but he decided that it was a new species.[28] Under the rules of botanical nomenclature, 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, Fawcett and Rendle linked Sloane's illustration to the species that was then known as F. suffocans Griseb. (a member of the subgenus Pharmacosycea). Gordon DeWolf agreed with their conclusion and used the name F. maxima for that species in the 1960 Flora of Panama.[29] Since this use has become widespread, Berg proposed that the name Ficus maxima be conserved in the way DeWolf had used it,[28] a proposal that was accepted by the nomenclatural committee.[30]
However, reassigning the name Ficus maxima did not leave F. aurea as the oldest name for this species. In 1822 Johann Heinrich Friedrich Link described Ficus ciliolosa. Berg concluded that the species Link described was actually F. aurea. Since Link's description predated Nuttall's by 24 years, priority should be given to the name F. ciliolosa. Since the name F. aurea is widely used and the name F. ciliolosa has not been, Berg proposed that the the name F. aurea be conserved.[28] 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 meant 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 is discovered.[30]
[edit] Synonyms
In 1920, Paul C. Standley described three new species based on collections from Panama and Costa Rica - Ficus tuerckheimii, F. isophlebia and F. jimenezii.[31] DeWolf concluded that they were all the same species,[29] and Berg synonymised them with F. aurea.[2] These names have been used widely for Mexican and Central American populations, and continue to be used by some authors.
[edit] References
- ^ a b Harvey, C. A.; W. A. Haber (1998). Remnant trees and the conservation of biodiversity in Costa Rican pastures. Agroforestry Systems 44 (1): 37-68. doi:.
- ^ a b c d e f Berg, C.C. (2007). Proposals for treating four species complexes in Ficus subgenus Urostigma section Americanae (Moraceae). Blumea 52: 295–312.
- ^ Simpson DP (1979). Cassell's Latin Dictionary, 5, London: Cassell Ltd., 883. ISBN 0-304-52257-0.
- ^ a b c d e f Bronstein, Judith L.; Aviva Patel (1992). Causes and Consequences of Within-Tree Phenological Patterns in the Florida Strangling Fig, Ficus aurea (Moraceae). American Journal of Botany 79 (1): 41-48. Botanical Society of America.
- ^ Flora de Nicaragua database. Tropicos. (Spanish)
- ^ Jousselin, Emmanuelle; Martine Hossaert-Mckey; Didier Vernet; Finn Kjellberg (2001). Egg deposition patterns of fig pollinating wasps: implications for studies on the stability of the mutualism. Ecological Entomology 26 (6): 602–608. doi:.
- ^ a b Janzen, Daniel H. (1979). How to be a fig. Annual Review of Ecology and Systematics 10: 13-51.
- ^ a b c d e f Bronstein, Judith L.; Martine Hossaert-McKey (1995). Hurricane Andrew and a Florida Fig Pollination Mutualism: Resilience of an Obligate Interaction. Biotropica 27 (3): 373-381. The Association for Tropical Biology and Conservation.
- ^ Putz, Francis E.; N. Michele Holbrook (1989). Strangler Fig Rooting Habits and Nutrient Relations in the Llanos of Venezuela. American Journal of Botany 76 (6): 781-788. Botanical Society of America.
- ^ a b c Swagel, Eric N.; A. Van H. Bernhard; George S. Ellmore (1997). Substrate water potential constraints on germination of the strangler fig Ficus aurea (Moraceae). American Journal of Botany 84 (5): 716-722. Botanical Society of America.
- ^ Horvitz, Carol C.; John B. Pascarella; Stephen McMann; Andrea Freedman; Ronald H. Hofstetter (1988). Functional Roles of Invasive Non-Indigenous Plants in Hurricane-Affected Subtropical Hardwood Forests. Ecological Applications 8 (4): 947-974. Ecological Society of America.
- ^ Elbert L. Little, Jr., Atlas of United States Trees Volume 5 Florida. Washington, D.C.: United States Government Printing Office, 1978
- ^ a b c Serrato, Alejandra; Guillermo Ibarra-Manríquez; Ken Oyama (2004). Biogeography and conservation of the genus Ficus (Moraceae) in Mexico. Journal of Biogeography 31 (3): 475-485. doi:.
- ^ Ficus aurea Nutt. Flora Mesoamericana: Lista Anotada. (Spanish)
- ^ Ficus aurea. Wunderlin, R. P., and B. F. Hansen. 2008. Atlas of Florida Vascular Plants. S. M. Landry and K. N. Campbell (application development), Florida Center for Community Design and Research. Institute for Systematic Botany, University of South Florida, Tampa.
- ^ Rutchey, K., T.N. Schall, R.F. Doren, A. Atkinson, M.S. Ross, D.T. Jones, M. Madden, L. Vilchek, K.A. Bradley, J.R. Snyder, J.N. Burch, T. Pernas, B. Witcher, M. Pyne, R. White, T.J. Smith III, J. Sadle, C.S. Smith, M.E. Patterson, and G.D. Gann. 2006. Vegetation Classification for South Florida Natural Areas. Saint Petersburg, Florida. United States Geological Survey, Open-File Report 2006-1240. 142 p.
- ^ Smith, Inge K.; John L. Vankat (1992). Dry Evergreen Forest (Coppice) Communities of North Andros Island, Bahamas. Bulletin of the Torrey Botanical Club 119 (2): 181-191. Torrey Botanical Society.
- ^ Howard, Richard A. (1950). Vegetation of the Bimini Island Group: Bahamas, B. W. I.. Ecological Monographs 20 (4): 317-349. Ecological Society of America.
- ^ Sillett, Stephen C.; S. Rob Gradstein; Dana Griffin, III (1995). Bryophyte Diversity of Ficus Tree Crowns from Cloud Forest and Pasture in Costa Rica. The Bryologist 98 (2): 251-260. American Bryological and Lichenological Society.
- ^ Wheelwright, Nathaniel T. (1985). Competition for dispersers, and the timing of flowering and fruiting in a guild of tropical trees. Oikos 44: 465-477.
- ^ Wheelwright, Nathaniel T. (1983). Fruits and the Ecology of Resplendent Quetzals. The Auk 100: 286-301.
- ^ Koptur, Suzanne (1992). Plants with Extrafloral Nectaries and Ants in Everglades Habitats. The Florida Entomologist 75 (1): 38-50. Florida Entomological Society.
- ^ a b c d Allen, Ginger M., Michael D. Bond, and Martin B. Main. 2002. 50 Common Native Plants Important In Florida's Ethnobotanical History. Circular 1439. University of Florida IFAS Extension.
- ^ Eldridge, Joan (October 1975). Bush medicine in the Exumas and Long Island, Bahamas. A field study. Economic Botany 29 (4): 307-332.
- ^ Adonizio, Allison L.; Kelsey Downum; Bradley C. Bennett; Kalai Mathee (2006). Anti-quorum sensing activity of medicinal plants in southern Florida. Journal of Ethnopharmacology 105 (3): 427-435. doi:.
- ^ Frodin, David G. (2004). History and concepts of big plant genera. Taxon 53 (3): 753–776.
- ^ Rønsted, N.; G.D. Weiblen; W.L. Clement; N.J.C. Zerega; V. Savolainen (2008). Reconstructing the phylogeny of figs (Ficus, Moraceae) to reveal the history of the fig pollination mutualism. Symbiosis 45 (1-3): 45-56.
- ^ a b c d Berg, Cornelis C. (May 2003). (1587–1590) Proposals to conserve the names Ficus citrifolia against F. caribaea, F. maxima with a conserved type, F. aurea against F. ciliolosa, and F. americana against F. perforata (Moraceae). Taxon 52: 368–370.
- ^ a b DeWolf, Gordon P., Jr. 1960. Ficus [Tourn.] L. In Lorin I. Nevling, Jr. Flora of Panama. Part IV. Fascicle II. Annals of the Missouri Botanical Garden, 47 (2):81-203
- ^ a b Brummitt, R.K. (May 2005). Report of the Committee for Spermatophyta: 56. Taxon 54 (2): 527-536.
- ^ Standley, Paul C. 1920. The Mexican and Central American Species of Ficus. Contributions from the United States National Herbarium 20(1):1-35

