TROPICAL FORESTS / Lecythidaceae 1745 mammals (e.g., squirrels and monkeys) and fish may Lecythidaceae also be important dispersers. S A Mori, New York Botanical Garden, New York, USA & 2004, Elsevier Ltd. All Rights Reserved. Economic Uses Historically, the Lauraceae is a very important economic family, especially the genus Cinnamomum, Introduction as a source of spices, e.g., cinnamon (C. verum), cassia bark (C. cassia) and laurel (Laurus nobilis), The Lecythidaceae is a pantropical family of trees perfume oils, e.g., rosewood oil (Aniba roseodora) found in the tropics of Central and South America, and sassafras oil (Ocotea odorifera), and pharma- southeast Asia, and Africa, including Madagascar ceuticals, e.g., camphor (C. camphora). However, the (Table 1). The family is divided into five subfamilies: most internationally important product produced (1) the Planchonioideae, with six genera, the best today is the avocado fruit (Persea americana). known of which is Barringtonia, and 59 species in Lauraceae wood is widely used locally, although a tropical Asia, Malaysia, northern Australia, the few are internationally important as high-quality Pacific Islands, and Madagascar; (2) the Foetidioi- timbers for furniture making or for resistance to salt deae, with a single genus, Foetidia, and 17 species in water, e.g., greenheart (Chlorocardium rodiaei), Madagascar, Mauritius, and East Africa; (3) the Borneo ironwood (Eusideroxylon zwageri), and Napoleonaeoideae, with two genera, Crateranthus Queensland walnut (Endiandra palmerstonii). Some and Naopoleonaea, and 11 species in West Africa; (4) species (e.g., Ocotea bullata), once internationally the Scytopetaloideae, with a single species, Aster- important, are now protected because of past over- anthos brasiliensis, in the Negro and Orinoco river exploitation. basins of Brazil and Venezuela and six genera and as many as 21 species in Africa; and (5) the Lecythi- See also: Ecology: Plant-Animal Interactions in Forest doideae of the western hemisphere. This article Ecosystems. Medicinal, Food and Aromatic Plants: focuses on the New World species of Lecythidaceae. Medicinal and Aromatic Plants: Ethnobotany and Con- New World Lecythidaceae range from Veracruz, servation Status. Tree Physiology: Physiology of Sexual Mexico (Eschweilera mexicana) to Paraguay (Car- Reproduction in Trees. Tropical Forests: Tropical Moist iniana estrellensis). The Caribbean harbors only Forests; Tropical Montane Forests. Grias cauliflora, a species found in Jamaica, Central America, and northwestern South America. Several Further Reading species of Eschweilera occur on Trinidad and Tobago, but these islands are South American, both Chanderbali AS, van der Werff H, and Renner SS geologically and in their floristic affinities. (2001) Phylogeny and historical biogeography of Laur- Neotropical Lecythidaceae are best known for the aceae: evidence from the chloroplast and nuclear Brazil nut of commerce (Bertholletia excelsa); for the genomes. Annals of the Missouri Botanical Garden 88: cannon-ball tree (Couroupita guianensis), which is 104–134. Kubitzki K and Kurz H (1984) Synchronized dichogamy planted as a botanical curiosity in tropical and and dioecy in neotropical Lauraceae. Plant Systematics subtropical botanical gardens; and because species and Evolution 147: 253–266. of the family are often ecological dominants in Renner SS (1999) Circumscription and phylogeny of the Amazonian forests. Laurales: evidence from molecular and morphological data. American Journal of Botany 86: 1301–1315. Taxonomy and Genetics Rohwer JG (1993) Lauraceae. In: Kubitski K, Rohwer JG, and Bittrich V (eds), The Families and Genera of In tropical America, there are 10 genera and 202 Flowering Plants, vol. 2, Flowering Plants: Dicotyledons known species of Lecythidoideae and a single species, – Magnoliid, Hamamelid and Caryophyllid families, pp. Asteranthos brasiliensis, of the African centered 366–391. Berlin, Germany: Springer-Verlag. Syctopetaloideae. The largest genus is Eschweilera Rohwer JG (2000) Toward a phylogenetic classification of with 85 species followed by Gustavia (41), Lecythis the Lauraceae: evidence from matK sequences. Systema- (26), Couratari (19), Cariniana (15), Grias (7), tic Botany 25: 60–71. Snow DW (1981) Tropical frugivorous birds and their food Corythophora (4), and Couroupita (3). Allantoma plants: a world survey. Biotropica 13: 1–14. lineata, Asteranthos brasiliensis,andBertholletia van der Werff H and Richter HG (1996) Toward an excelsa belong to monotypic genera. improved classification of Lauraceae. Annals of the Molecular studies have demonstrated that the Missouri Botanical Garden 83: 409–418. Lecythidaceae are monophyletic if members of the 1746 TROPICAL FORESTS / Lecythidaceae Table 1 Subfamilies and genera of Lecythidaceae according to Morton et al. (1998) Subfamily Region Genera Number of species FOETIDIOIDEAE Madagascar, Mauritius, and east African tropics Foetidia 17 LECYTHIDOIDEAE New World tropics Allantoma 1 Bertholletia 1 Cariniana 16 Corythophora 3 Couratari 19 Couroupita 3 Eschweilera 85 Grias 7 Gustavia 41 Lecythis 26 NAPOLEONAEOIDEAE African tropics Crateranthus 3 Napoleonaea 8 PLANCHONIOIDEAE African and Asian tropics Abdulmajidia 2 Barringtonia 41 Petersianthus 2 Chydenanthus 2 Careya 4 Planchonia 8 SCYTOPETALOIDEAE African tropics and Asteranthos in Amazonia Asteranthos 1 Brazzeia 3 Oubanguia 3 Pierrina 1 Rhaptopetalum 10 Scytopetalum 3 Scytopetalaceae are included. The relationship of nian forests. In this plot, there are 7791 trees of Asteranthos brasiliensis with the Scytopetalaceae is Lecythidaceae equal to or greater than 10 cm in supported by similarities in embryology, the presence diameter at 1.37 m (dbh, diameter at breast height). in both of ruminate endosperm, and molecular Among these trees are 38 different species, or nearly affinities. 19% of all of the species of Lecythidaceae known Ongoing molecular studies of neotropical Lecythi- from the Neotropics. In each hectare, there are 45 to daceae indicate that not all of the genera are 149 individuals and 11 to 24 species. In this forest, monophyletic. Cariniana, Eschweilera, and Lecythis, where there are about 285 species of trees in all for example, include several clades that may have to families in this size class per hectare, the Brazil nut be segregated as separate genera. Other genera, for family accounts for 12% of the individuals and 6% example Corythophora, Couratari, Couroupita, of the species of trees. Grias,andGustavia, form natural groups as they Species of the Brazil nut family diminish in are currently defined. importance at elevations above 1000 m, in periodi- cally flooded forests, and in extremely dry habitats such as the llanos of Colombia and Venezuela and Ecology the caatinga of northeastern Brazil. New World Lecythidaceae are often dominant in Nearly all neotropical Lecythidaceae are trees of lowland forests on well-drained soils between 191 N either the understory, canopy, or emergent layers. and 251 S latitudes. They rank as the first or second The tallest, for example the forest-dwelling Carini- most important family of trees in frequency, density, ana micrantha and Couratari stellata, may reach and dominance in the vicinity of Bele´m, Brazil; the 55–60 m in height. These genera, both dispersed by third most important at La Fume´e Mountain, French the wind, possess species of smaller stature when Guiana; and one of the most important in central they are found in more open habitats; for example, Amazonian Brazil. Couratari pyramidata of the cerrado of central A 100-ha plot established for the study of Brazil. The smallest is Eschweilera nana, a species Lecythidaceae as part of the Biological Dynamics of that often possesses an underground trunk adapting Forest Fragments Project in central Amazonian it to survive the frequent fires of the cerrado. Brazil about 80 km north of Manaus illustrates the Individuals of E. nana may only reach several meters ecological importance of Lecythidaceae in Amazo- in height. TROPICAL FORESTS / Lecythidaceae 1747 Trees of Bertholletia excelsa in the 140–150 cm dbh Pollination Biology size class have been estimated by radiocarbon dating to be only 270 years old. One of the oldest known The Brazil nut family provides examples of plant– trees from the Neotropics is an individual of Cariniana animal coevolution. The flowers, fruits, and seeds are micrantha radiocarbon dated at 1400 years old. morphologically and anatomically adapted to the Species of Lecythidaceae are not common in animals that pollinate their flowers and the animals, secondary vegetation because they are extremely wind, and water that disperse their seeds. vulnerable to fire. If a forest is cut down and not Adaptation of Lecythidaceae for pollination has subjected to fire, many Lecythidaceae will sprout taken place, for the most part, in the male part of from the cut stumps and flower and set fruit from the flower. There are two types of flowers, radially the new branches. However, if fire follows cutting and bilaterally symmetrical and three different most Lecythidaceae are killed; hence old slash-and- rewards offered to pollinators – fertile pollen, sterile burn fields throughout the Neotropics harbor few pollen, and nectar (Figure 1). Radially symmetrical species and individuals of Lecythidaceae. Species flowers, such as those of Gustavia superba (Figure 2), of Lecythidaceae are generally indicators of undis- offer only fertile pollen as a reward; consequently turbed habitats. the pollinator reward and the pollen