TROPICAL ECOSYSTEMS / Ficus spp. (and other important Moraceae) 1699 breaches of the NVC increasing steadily to more than House SM (1997) Reproductive biology of eucalypts. In: 200 per year, with only seven prosecutions com- Williams JE and Woinarski JCZ (eds) Eucalypt Ecology: menced out of more than 700 alleged breaches from Individuals to Ecosystems, pp. 30–55. Cambridge, UK: 1998 to April 2002. Cambridge University Press. There are many implications of the changes to the Keith H (1997) Nutrient cycling in eucalypt ecosystems. In: eucalypt forests across Australia. The clearing of Williams JE and Woinarski JCZ (eds) Eucalypt Ecology: Individuals to Ecosystems, pp. 197–226. Cambridge, native habitat is the single most threatening process UK: Cambridge University Press. for biodiversity loss and species extinction in Kirkpatrick JB (1997) Vascular plant–eucalypt interac- Australia. The loss of species in Australia over the tions. In: Williams JE and Woinarski JCZ (eds) Eucalypt past two centuries due to human impact is con- Ecology: Individuals to Ecosystems, pp. 227–245. Cam- servatively estimated at 97 plants, 17 species of bridge, UK: Cambridge University Press. mammal, and three birds. However, several hundred Landsberg JJ and Cork SJ (1997) Herbivory: Interactions vertebrates, several thousand plants, and an untold between eucalypts and the vertebrates and invertebrates number of invertebrates are threatened with extinc- that feed on them. In: Williams JE and Woinarski JCZ tion due to loss of habitat. The extent and rate (eds) Eucalypt Ecology: Individuals to Ecosystems, pp. of deforestation in Queensland are comparable to 342–372. Cambridge, UK: Cambridge University Press. that in Western Australia and Victoria in the May TW and Simpson JA (1997) Fungal diversity and ecology in eucalypt ecosystems. In: Williams JE and past, which has given rise to significant salinity Woinarski JCZ (eds) Eucalypt Ecology: Individuals to problems in these states, both in terms of dryland Ecosystems, pp. 246–277. Cambridge, UK: Cambridge salinity and salinization of drinking and irrigation University Press. water. The release of carbon into the atmosphere by National Forest Inventory website. www.affa.gov.au/con- clearing operations also contributes significantly tent/output.cfm?&CONTTYPE ¼ outputs&OBJECTID (13% in 1996) to Australia’s greenhouse-gas emis- ¼ D52EDDBA-23DF-485A-B1C968D1C08E79D1. sion, and there is a net loss of carbon dioxide equivalent to a ratio of 3 : 1 compared with that captured by vegetation acting as a sink. Although many of these consequences of the loss of eucalypt Ficus spp. (and other important forests have been known for decades, and have been recognized by governments and addressed in Moraceae) recent policy initiatives, clearing of eucalypt forests J K Francis, Jardı´n Bota´nico Sur, San Juan, Puerto continues. Rico, USA & 2004, Elsevier Ltd. All Rights Reserved. See also: Ecology: Plant-Animal Interactions in Forest Ecosystems; Reproductive Ecology of Forest Trees. Entomology: Bark Beetles; Defoliators; Foliage Feeders in Temperate and Boreal Forests; Sapsuckers. Pathol- Introduction ogy: Diseases of Forest Trees; Insect Associated Tree Moraceae is a family of mostly tropical trees Diseases. Tree Breeding, Practices: Genetic Improve- ment of Eucalypts; Mycorrhizae. containing 37 genera and about 1100 species. The most important genus, both in terms of numbers and benefit to humans, is Ficus. The family is character- ized as having milky or sometimes watery latex. Further Reading Leaves are mostly alternate, simple, entire, and Fox MD (1999) Present environmental influences on the pinnately veined. The flowers are unisexual but Australian flora. In: Orchard AE and Thompson HS (eds) variable in form, as are the inflorescences. The fruits Flora of Australia, 2nd edn, vol. 1, pp. 205–249. are commonly drupaceous, embedded in a fleshy Melbourne, Australia: ABRS/CSIRO Australia. receptacle forming a syncarp. The seeds are large Gill AM (1997) Eucalypts and fires: interdependent or without endosperm or small with endosperm. independent? In: Williams JE and Woinarski JCZ (eds) Although a few of the species produce timber, bark, Eucalypt Ecology: Individuals to Ecosystems, pp. 151– and yield medicinal products, the greatest benefit to 167. Cambridge, UK: Cambridge University Press. humans and animals is from the fruits that many Groves RH (ed.) (1997) Australian Vegetation, 2nd edn. Cambridge, UK: Cambridge University Press. produce. Except for a few of the fruit-producing Groves RH (1999) Present vegetation types. In: Orchard species, three or four timber species, and a number of AE and Thompson HS (eds) Flora of Australia, 2nd edn, Ficus and other species planted for ornament and vol. 1, pp. 369–402. Melbourne, Australia: ABRS/ shade, members of the family are little planted or CSIRO Australia. actively managed. 1700 TROPICAL ECOSYSTEMS / Ficus spp. (and other important Moraceae) Taxonomy/Genetics The family Moraceae consists of trees with lesser numbers of shrubs, climbers, and herbs, most of Note: The following treatment is based on taxonomy which are tropical. All (except for the genus Fatoua) that predates the publication of the Angiosperm secrete milky (sometimes watery) latex from laticifers Phylogeny Group’s higher-level classification of the in parenchymatous tissues in stems and sometimes angiosperms. Moraceae (used here in the narrow leaves. Leaves are alternate or rarely subopposite, sense) is one of the ewosid families of Rosales. most frequently simple, entire, and pinnately veined. Moraceae is part of the order Urticales, which is Stipules are present, although often fused and forming believed to have descended from a single ancestral a cap over the bud. Inflorescences take various forms. line. Because of morphological and molecular Flowers are unisexual, often four-merous. Plants may similarity, it has been suggested that Moraceae, be monoecious or dioecious. Although some species Celtidaceae, Cecropiaceae, and Urticaceae should have dry fruits, most have drupaceous or achene fruits be merged. The genera of Cecropiaceae appear under embedded in fleshy receptacles. The family is further Moraceae in older references. Moraceae currently divided into the tribes Moreae (Morus, Broussonetia, contains 37 genera and about 1100 species (Table 1). Milicia, Maclura, Trophis, Streblus, Bleekrodea,and Fatoua), which have ‘urticaceous’ stamens; tribe Artocarpeae (Artocarpus, Paratocarpus, Triculia, Table 1 Details of genera and species within Moraceae Prainea, Hullettia, Antiaropsis, Sparattosyce, Bato- Genus No. sp. Range Life forms carpus, Bagassa, Sorocea, Clarisia,andPoulsenia), Antiaris 1 Africa, Australasia t with generally unisexual inflorescences and seeds Antiaropsis 1 New Guinea t, s almost without endosperm; tribe Castilleae (Perebea, Artocarpus B50 Asia, Australia t Maquira, Castilla, Helicostylis, Pseudolmedia, Nau- Bagassa 1 America t cleopsis, Antiaris,andMesogyne), with spontaneous Batocarpus 4 America t abscission of branches; tribe Dorstenieae (Utsetela, Bleekrodea 3 Melesia, Madagascar t, s Bosqueiopsis 1 Africa t, s Bosqueiopsis, Helianthostylis, Trymatococcus, Brosi- Brosimum 13 America t mum, Trilepisium, Scyposyce,andDorstenia)usually Broussonetia 8 Asia, Madagascar t, s, c with uncinate hairs; and tribe Ficeae (Ficus), which Castilla 3 America t have syconia enclosing the flowers. Clarisia 3 America t, s The family follows various growth models. In Dorstenia B105 Pan tropical s, h Fatoua 3 Pacific, Madagascar s, h Troll’s model the architecture is built by the continual Ficus B750 Pan tropical t, s, c superposition of plagiotropic branches, each division Helicostylis 7 America t alternating between main-line axis and determinate Helianthostylis 2 America t branch, and is followed by Milicia excelsa, in Rauh’s Hullettia 2 Malaya t, s model, a monopodial trunk grows rhythmically and Maclura 11 North America, Pan t, s, c tropical develops tiers of branches and the branches are Maquira 5 America t morphologically identical to the trunk, is exemplified Mesogyne 1 Africa t, s by Artocarpus heterophyllus, Ficus aurea,andMu- Milicia 2 Africa t sanga cercropoides. The architecture in Cook’s model Morus 10–15 Almost worldwide t results when a monopodial trunk with spiral or Naucleopsis 20–25 America t Parartocarpus 3 Malesia, Pacific t decussate phyllotaxis adds branches phylomorphi- Perebea 9–10 America t, s cally. Castilla follows this model. Roux’s model, Poulsenia 1–2 America t followed by Antiaris welwitschii, Milicia regia,and Prainea 4 Malaysia-New Guinea t, c Perebea guianensis, occurs when a monopodial Pseudolmedia 8–9 America t orthotropic trunk meristem grows continuously and Scyphosyce 2 Africa s Sorocea B20 America t, s has plagiotropic branches that are inserted continu- Sparattosyce 1 New Caledonia t ously. Dorstenia and F. theophrastoides grow by Streblus B25 Old World t, s Corner’s model in which vegetative growth of a single Treculia 3 Africa, Madagascar t, s aerial meristem produces one unbranched axis with Trilepisium 1 Africa, Indian Ocean t lateral inflorescences. Trophis 9 America, Asia t, s Trymatococcus 3 America t Sex determination in F. carica, and probably other Utsetela 1 Africa s members of the subgenus Ficus, is different from the XX/XY method used in most other life forms, t, trees; s, shrubs; c, climbers; h, herbs. Information source: Kubitzki K, Rohwer JG, and Bittrich V (1993) from mosses through humans. It is believed to be The Families and Genera of Vascular Plants, vol. 2.