Ficus macrophylla

Moreton Bay fig
A specimen at The Domain, Sydney, planted in 1850
Scientific classification
Kingdom: Plantae
(unranked): Angiosperms
(unranked): Eudicots
(unranked): Rosids
Order: Rosales
Family: Moraceae
Genus: Ficus
Species: F. macrophylla
Binomial name
Ficus macrophylla
Desf. ex Pers.
Synonyms

Ficus macrocarpa Hugel ex Kunth & Bouche
Ficus magnolioides[1]

Ficus macrophylla, commonly known as the Moreton Bay fig or Australian banyan, is a large evergreen banyan tree of the family Moraceae that is a native of most of the eastern coast of Australia, from the Atherton Tableland (17° S) in the north to the Illawarra (34° S) in New South Wales, and Lord Howe Island. Its common name is derived from Moreton Bay in Queensland, Australia. It is best known for its beautiful buttress roots.

As Ficus macrophylla is a strangler fig, seed germination usually takes place in the canopy of a host tree and the seedling lives as an epiphyte until its roots establish contact with the ground. It then enlarges and strangles its host, eventually becoming a freestanding tree by itself. Individuals may reach 60 m (200 ft) in height. Like all figs, it has an obligate mutualism with fig wasps; figs are only pollinated by fig wasps, and fig wasps can only reproduce in fig flowers.

Ficus macrophylla is widely used as a feature tree in public parks and gardens in warmer climates such as California, Portugal, Italy (Sicily, Sardinia and Liguria), northern New Zealand (Auckland), and Australia. Old specimens can reach tremendous size. Its aggressive root system allows its use in only the largest private gardens.

Taxonomy

Christiaan Hendrik Persoon published a description of the Moreton Bay fig in his 1807 Synopsis Plantarum, the material having been described by French botanist René Louiche Desfontaines. The type specimen has been lost but was possibly located in Florence.[2] The specific epithet macrophylla is derived from the Ancient Greek makro "large" and phyllon "leaf",[3] and refers to the size of the leaves.[4]

Charles Moore described Ficus columnaris in 1870 from material collected from Lord Howe Island, choosing the species name from the Latin columnaris for the column-like roots. English botanist E. J. H. Corner reduced this to synonymy with F. macrophylla in 1965, before P.S. Green noted it was distinct enough in its morphology for subspecies status in 1986. Australian botanist Dale J. Dixon reviewed material and felt the differences too minor to warrant subspecific status,[2] and recognised two forms: Ficus macrophylla f. macrophylla, a free-standing tree which is endemic to mainland Australia and Ficus macrophylla f. columnaris, a hemiepiphyte without a distinct main trunk, which is endemic to Lord Howe Island.[5] The species is generally commonly known as the Moreton Bay fig,[4] after Moreton Bay in southern Queensland, although it is found elsewhere. The term has also been mistakenly generalised to other fig species in Australia. An alternate name—black fig—is derived from the dark colour of the ageing bark.[6]

With over 750 species, Ficus is one of the largest angiosperm genera.[7] Based on morphology, Corner divided the genus into four subgenera;[5] later expanded to six.[8] In this classification, the Moreton Bay fig was placed in subseries Malvanthereae, series Malvanthereae, section Malvanthera of the subgenus Urostigma.[9] In his reclassification of the Australian Malvanthera, Dixon altered the delimitations of the series within the section, but left this species in series Malvanthereae.[5]

In 2005, Cornelis Berg completed Corner's treatment of the Moraceae for the Flora Malesiana; the completion of that work had been delayed since 1972 as a result of disagreements between Corner and C. J. J. G. van Steenis, editor of the Flora Malesiana.[10] Berg combined sections Stilpnophyllum and Malvanthera into an expanded section Stilpnophyllum. This left the Moreton Bay fig in subsection Malvanthera, section Stilpnophyllum.[9]

Based on DNA sequences from the nuclear ribosomal internal and external transcribed spacers, Nina Rønsted and colleagues rejected previous subdivisions of the Malvanthera. Instead, they divided section Malvanthera into three subsections—Malvantherae, Platypodeae and Hesperidiiformes. In this system, the Moreton Bay fig is in the new subsection Malvantherae, along with F. pleurocarpa. The Malvantherae appear to be an early offshoot and basal to the group. Both forms are native to Australia, with form columnaris of macrophylla having colonised Lord Howe Island. Ronsted and colleagues suggest this last form might be the most ancient of the three taxa.[9] The section Malvanthera itself is thought to have evolved 41 million years ago and radiated from 35 million years ago.[8]

Description

The Moreton Bay fig is an evergreen tree that can reach heights of 60 m (200 ft).[1] The trunk can be massive, with thick, prominent buttressing, and reach a diameter of 2.4 m (8 ft).[4] The rough bark is grey-brown,[11] and marked with various blemishes.[12] It is monoecious: each tree bears functional male and female flowers.[5] As implied by its specific epithet, it has large, elliptic, leathery, dark green leaves, 15–30 cm (6–12 in) long, and they are arranged alternately on the stems. The leaves and branches bleed a milky sap if cut or broken. The figs are 2–2.5 cm (0.75–1 in) in diameter, turning from green to purple with lighter spots as they ripen;[1] ripe fruit may be found year round,[12] although more abundant from February to May.[4] Although edible, they are unpalatable and dry.[4]

The characteristic "melting" appearance of the Moreton Bay fig is due to its habit of dropping aerial roots from its branches, which upon reaching the ground, thicken into supplementary trunks which help to support the weight of its crown.

It is a rainforest plant and in this environment more often grows in the form of an epiphytic strangler vine than that of a tree. When its seeds land in the branch of a host tree it sends aerial, "strangler" roots down the host trunk, eventually killing the host and standing alone.[4]

Reproduction

Pleistodontes frogatti, the fig wasp that pollinates the Moreton Bay fig

Figs have an obligate mutualism with fig wasps (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. The mainland and Lord Howe populations of the Moreton Bay fig are both pollinated by the fig wasp Pleistodontes froggatti.[2][13]

As is the case with all figs, the fruit is actually an inverted inflorescence known as a syconium, with tiny flowers arising from the inner surface.[12] Ficus macrophylla is monoecious—both male and female flowers are found on the same plant, and in fact in the same fruit although they mature at different times. Female wasps enter the syconium and lay eggs in the female flowers as they mature. These eggs later hatch and the progeny mate. The females of the new generation collect pollen from the male flowers, which have matured by this point, and leave to visit other syconia and repeat the process. A field study in Brisbane found that a F. macrophylla trees often bore both male and female syconia at the same time—which could be beneficial for reproduction in isolated populations such as those on islands. The same study found that male phase syconia development persisted through the winter, showing that its wasp pollinator tolerated cooler weather than those of more tropical fig species. F. macrophylla itself can endure cooler climates than other fig species.[14]

Distribution and habitat

Emergent Moreton Bay fig in situ, estimated 50 metres tall, Davis Scrub Nature Reserve, Australia.

The Moreton Bay fig is a native of most of the eastern coast, from the Atherton Tableland (17° S) in north Queensland, to the Shoalhaven River (34° S) on New South Wales south coast.[12] It is found in subtropical, warm temperate and dry rainforest,[15] where, as an emergent tree, its crown may tower above the canopy,[12] particularly along watercourses on alluvial soils.[16] It often grows with trees such as white booyong (Argyrodendron trifoliolatum), Flindersia species, giant stinging tree (Dendrocnide excelsa), lacebark (Brachychiton discolor), red cedar (Toona ciliata), hoop pine (Araucaria cunninghamii), green-leaved fig (Ficus watkinsiana) and Cryptocarya obovata.[15] The soils it grows on are high in nutrients, and include Bumbo Latite and Budgong Sandstone.[16] As rainforests were cleared, isolated specimens were left standing in fields as remnant trees,[17] valued for their shade and shelter for livestock.[18] One such tree was a landmark for and gave its name to the Wollongong suburb of Figtree in New South Wales.[17]

Ecology

The huge numbers of fruit produced by the Moreton Bay fig make it a key source of food in the rainforest.[18] It is an important food to the fruit-eating pigeons such as the wompoo fruit-dove (Ptilinopus magnificus), and topknot pigeon (Lopholaimus antarcticus), and a sometime food of the rose-crowned fruit-dove (Ptilinopus regina).[19] Other bird species include the yellow-eyed cuckoo-shrike (Coracina lineata), pied currawong (Strepera graculina), Australasian figbird (Sphecotheres vieilloti), green catbird (Ailuroedus crassirostris), Regent bowerbird (Sericulus chrysocephalus), satin bowerbird (Ptilonorhynchus violaceus), and Lewin's honeyeater (Meliphaga lewinii).[4] Fruit bats such as the grey-headed flying-fox (Pteropus poliocephalus) also feed on the fruit.[20] In addition to the pollinating wasp, Pleistodontes froggatti, syconia of the Moreton Bay fig are host to several species of non-pollinating chalcidoid wasps including Sycoscapter australis (Pteromalidae), Eukobelea hallami (Sycophaginae) and Meselatus sp. (Epichrysomallinae).[21]

The thrips species Gynaikothrips australis feeds on the underside of new leaves of F. macrophylla, as well as F. rubiginosa and F. obliqua. As plant cells die, nearby cells are induced into forming meristem tissue and a gall results, and the leaves become distorted and curl over. The thrips begin feeding when the tree has flushes of new growth, and the life cycle is around six weeks. At other times, thrips reside on old leaves without feeding. The species pupates sheltered in the bark. The thrips remain in the galls at night and wander about in the daytime and return in the evening, possibly to different galls about the tree.[22]

Stressed trees can also be attacked by psyllids to the point of defoliation. Grubs hatch from eggs laid on the edges of leaves and burrow into the leaf to suck nutrients, the tree's own latex shielding the insect.[16] Caterpillars of the moth species Eustixis caminaea can strip trees of their leaves.[16] The tree is also a host for the longhorn beetle species Agrianome spinicollis.[16]

Potentially invasive species

A young Moreton Bay fig starts life as an epiphyte in an Auckland, New Zealand park

Ficus macrophylla is commonly cultivated in Hawaii and in northern New Zealand. In both places, it has now naturalised, having acquired its pollinating wasp (Pleistodontes froggatti). In Hawaii the wasp was deliberately introduced in 1921, and in New Zealand it was first recorded in 1993, having apparently arrived by long-distance dispersal from Australia. The arrival of the wasp led to prolific production of fruits containing many small seeds adapted for dispersal by birds. The Moreton Bay fig has been found growing on both native and introduced trees in New Zealand and in Hawaii. The size and vigour of this fig in New Zealand, and its lack of natural enemies, as well as its immunity to possum browsing, indicate that it may be able to invade forest and other native plant communities.[1][23]

Uses

The Moreton Bay fig has been widely used in public parks in frost-free areas, and was popular with early settlers of Australia.[11] Around the beginning of the 20th century, the director of the Royal Botanic Gardens in Sydney, Joseph Maiden, advocated the planting of street trees, generally uniform rows of the one species. He recommended Moreton Bay figs be spaced at 30 m (100 ft) intervals—far enough to avoid crowding as the trees matured but close enough so that their branches would eventually interlock.[24] Specimens can reach massive proportions, and have thrived in drier climates; impressive specimen trees have been grown in the Waring Gardens in Deniliquin, and in Hay.[25] They can withstand light frosts and can cope with salt-laden spray in coastal situations, and their fruit is beneficial for urban wildlife. However their huge size precludes use in gardens, and their roots are highly invasive and can damage piping and disrupt footpaths and roadways. The vast quantities of crushed fruit can be messy on the ground.[18]

Although their root buttressing is a potential feature, the Moreton Bay fig is poorly suited to bonsai as their large leaves do not reduce much in size and they have long internodal growth.[26] It is can be used as an indoor plant in medium to brightly lit indoor spaces.[27]

The soft light timber has a wavy texture and is used for cases.[4] Aborigines traditionally used the fibres for fishing nets.[4]

Notable specimens

Trunk of F. macrophylla at Orto botanico di Palermo, showing buttressed roots

Large specimens of Moreton Bay fig trees are found in many parks and properties throughout eastern and north-eastern Australia. The Brisbane, Melbourne and Sydney botanical gardens contain numerous specimens planted in the middle of the nineteenth century. These trees are, on average, 30 to 35 metres tall and 40 metres wide. Currently the tallest Moreton Bay fig is found on a cattle farm near Egg Rock in the Numinbah valley of Queensland and measured 49 metres (161 ft) in 2006. At Mount Keira, near Wollongong there is a Moreton Bay fig measured at 58 metres (190 ft) tall.[28] There are many large specimens in New Zealand. A Moreton Bay fig at Pahi on the Kaipara Harbour, Northland, was measured in 1984 as 26.5 metres (87 ft) high and 48.5 metres (159 ft) wide.[29]

Several large specimens grow in California, United States. The tallest Ficus macrophylla in North America is adjacent to San Diego's Natural History Museum and was planted in 1914. By 1996 it stood 23.7 metres (78 ft) high and 37.4 metres (123 ft) across.[30] The widest Moreton Bay fig in North America is Santa Barbara's Moreton Bay Fig Tree. It was planted in 1876, reportedly by a little girl who was given a seedling by an Australian sailor.[31] It measured 53.6 metres (176 ft) across in 1997. It is still a popular Santa Barbara tourist attraction.[32] Santa Monica resident and US Senator John P. Jones planted a Moreton Bay fig in a corner of his estate in 1889, which became the patio of the Miramar Hotel in 1921.[33] Another large Los Angeles Moreton Bay fig, known as the Aoyama Tree, stands between the Japanese American National Museum and the Temporary Contemporary downtown. It was planted by Buddhist Japanese Americans in the early 20th century.[34]

There is a notable specimen sprawling on steps at the Botanical Garden of the University of Coimbra, Portugal.[35] Impressive specimens also grow in several parks in Liguria (Bordighera and Sanremo) and in the Orto Botanico in Palermo, Italy, and in some squares in the oldest part of the city.

Two South African specimens, in the Arderne Gardens in Claremont and the Pretoria Zoo respectively, have the widest and second widest canopies of any single-stemmed trees in the country. The Pretoria specimen was planted before 1899, and was 27 metres (89 ft) tall with a canopy width of 43.1 metres (141 ft) by 2012.[36][37]

References

  1. 1 2 3 4 Starr F, Starr K, and Loope L. Ficus macrophylla - Moreton bay fig - Moraceae United States Geological Survey--Biological Resources Division Haleakala Field Station, Maui, Hawai'i, 2003.
  2. 1 2 3 Dixon, Dale J. (2001). "Figs, wasps and species concepts: a re-evaluation of the infraspecific taxa of Ficus macrophylla (Moraceae: Urostigma sect. Malvanthera)". Australian Systematic Botany 14 (1): 125–132. doi:10.1071/SB99026.
  3. ↑ Liddell, Henry George and Robert Scott (1980). A Greek-English Lexicon (Abridged Edition). United Kingdom: Oxford University Press. ISBN 0-19-910207-4.
  4. 1 2 3 4 5 6 7 8 9 Floyd, Alex G. (2009). Rainforest Trees of Mainland Southeastern Australia. Lismore, NSW: Terania Rainforest Publishing. pp. 231–232. ISBN 978-0-9589436-7-3.
  5. 1 2 3 4 Dixon, Dale J. (2003). "A taxonomic revision of the Australian Ficus species in the section Malvanthera (Ficus subg. Urostigma: Moraceae)" (pdf). Telopea 10 (1): 125–153.
  6. ↑ Webber, Len (1991). Rainforest to Bonsai. East Roseville, NSW: Simon and Schuster. p. 105. ISBN 0-7318-0237-3.
  7. ↑ Frodin, David G. (2004). "History and concepts of big plant genera". Taxon 53 (3): 753–776. doi:10.2307/4135449. JSTOR 4135449.
  8. 1 2 Rønsted, Nina; Weiblen, George D.; Clement, W. L.; Zerega, N. J. C.; Savolainen, V. (2008). "Reconstructing the phylogeny of figs (Ficus, Moraceae) to reveal the history of the fig pollination mutualism" (PDF). Symbiosis 45 (1–3): 45–56.
  9. 1 2 3 Rønsted, Nina; Weiblen, George D.; Savolainen, V; Cook, James M. (2008). "Phylogeny, biogeography, and ecology of Ficus section Malvanthera (Moraceae)" (PDF). Molecular Phylogenetics and Evolution 48 (1): 12–22. doi:10.1016/j.ympev.2008.04.005. PMID 18490180.
  10. ↑ Weiblen, G. D.; W. L. Clement (2007). "Flora Malesiana. Series I. Volume 17 parts 1 & 2" (PDF). Edinburgh Journal of Botany 64 (3): 431–437. doi:10.1017/S0960428607064311.
  11. 1 2 Holliday, Ivan (1989). A Field Guide to Australian Trees. Melbourne: Hamlyn Australia. p. 198. ISBN 0-947334-08-4.
  12. 1 2 3 4 5 Fairley A, Moore P (2000). Native Plants of the Sydney District:An Identification Guide (2nd ed.). Kenthurst, NSW: Kangaroo Press. p. 62. ISBN 0-7318-1031-7.
  13. ↑ Lopez-Vaamonde, Carlos; Dale J. Dixon; James M. Cook; Jean-Yves Rasplus (2002). "Revision of the Australian species of Pleistodontes (Hymenoptera: Agaonidae) fig-pollinating wasps and their host-plant associations". Zoological Journal of the Linnean Society 136 (4): 637–683. doi:10.1046/j.1096-3642.2002.00040.x.
  14. ↑ McPherson, John R. (2005). "Phenology of Six Ficus L., Moraceae, Species and its Effects on Pollinator Survival, in Brisbane, Queensland, Australia". Geographical Research 43 (3): 297–305. doi:10.1111/j.1745-5871.2005.00329.x.
  15. 1 2 Boland, Douglas J.; Brooker, M. I. H.; Chippendale, G. M.; McDonald, Maurice William (2006). Forest trees of Australia. Collingwood, Victoria: CSIRO Publishing. pp. 193–195. ISBN 0-643-06969-0.
  16. 1 2 3 4 5 Benson, Doug; McDougall, Lyn (1997). "Ecology of Sydney Plant Species Part 5: Dicotyledon Families Flacourtiaceae to Myrsinaceae" (PDF). Cunninghamia 5 (2): 330–544.
  17. 1 2 Fuller, Leon (1980). Wollongong's Native Trees. Kiama, New South Wales: Weston & Co. pp. 218–219. ISBN 0-9594711-0-3.
  18. 1 2 3 Elliot, Rodger W.; Jones, David L.; Blake, Trevor (1986). Encyclopaedia of Australian Plants Suitable for Cultivation: Volume 4 (Eu-Go). Port Melbourne, Victoria: Lothian Press. pp. 280, 286. ISBN 0-85091-589-9.
  19. ↑ Innis, Gary John (1989). "Feeding Ecology of Fruit Pigeons in Subtropical Rainforests of Southeast Queensland". Aust. Wildl. Res. 16 (4): 365–394. doi:10.1071/WR9890365.
  20. ↑ Eby., P. "Diet Species of the Grey-headed Flying-fox in the Sydney Region". Ku-ring-gai Bat Conservation Society Inc. Retrieved 3 May 2009.
  21. ↑ Bean, Daniel; Cook, James M. (2001). "Male mating tactics and lethal combat in the nonpollinating fig wasp Sycoscapter australis". Animal Behaviour 62 (3): 535–542. doi:10.1006/anbe.2001.1779.
  22. ↑ Tree, Desley J; Walter, G. H. (2009). "Diversity of host plant relationships and leaf galling behaviours within a small genus of thrips –Gynaikothrips and Ficus in south east Queensland, Australia". Australian Journal of Entomology 48 (4): 269–275. doi:10.1111/j.1440-6055.2009.00706.x.
  23. ↑ Gardner, Rhys O.; John W. Early (1996). "The naturalisation of banyan figs (Ficus spp., Moraceae) and their pollinating wasps (Hymenoptera: Agaonidae) in New Zealand". New Zealand Journal of Botany 34: 103–110. doi:10.1080/0028825x.1996.10412697.
  24. ↑ Frawley, Jodi (2009). "Campaigning for Street Trees, Sydney Botanic Gardens 1890s-1920s" (PDF). Environment and History 15 (3): 303–322. doi:10.3197/096734009x12474738199953.
  25. ↑ de Beuzeville WAW (1947). Australian Trees for Australian Planting. Sydney: Forestry Commission of New South Wales/ A. H. Pettifer, Government Printer. pp. 47–48.
  26. ↑ Koreshoff, Dorothy and Vita (1984). Bonsai with Australian native Plants. Brisbane: Boolarong Publications. p. 52. ISBN 0-908175-66-3.
  27. ↑ Ratcliffe, David & Patricia (1987). Australian Native Plants for Indoors. Crows Nest, NSW: Little Hills Press. p. 90. ISBN 0-949773-49-2.
  28. ↑ "Moreton Bay Fig – Mount Keira". National Tree Register of Big Trees. Retrieved 23 November 2011.
  29. ↑ S.W. Burstall and E.V. Sale (1984). Great Trees of New Zealand. Wellington: Reed. p. 39. ISBN 0-589-01532-X.
  30. ↑ "Ficus macrophylla: Moreton Bay Fig" SDNHM Botany Department.
  31. ↑ Baker, Gayle (2003). Santa Barbara. Santa Barbara, CA: HarborTown Histories. p. 55. ISBN 9780971098411.
  32. ↑ The Santa Barbara Fig Tree
  33. ↑ 20. Miramar Moreton Bay Fig Tree Santa Monica Landmarks
  34. ↑ The Cultural Landscape Foundation (2010). "Aoyama Tree". Every Tree Tells a Story: The Cultural Landscape Foundation's 2010 Landslide. Washington, D.C.: The Cultural Landscape Foundation. Retrieved 10 January 2012.
  35. ↑ Pakenham, Thomas (2002). Remarkable Trees of the World. London: Weidenfeld & Nicolson. pp. 146–147. ISBN 0-297-84300-1.
  36. ↑ Versluis, Jeanne-Marié (4 January 2012). "Reuse-vyeboom op kampioen-lys". Beeld. Retrieved 9 January 2013.
  37. ↑ Schwan, Angeliné. "Who planted this massive tree?". National Zoological Gardens of South Africa. National Research Foundation. Retrieved 11 December 2015.

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