Palaeobolalli.,/ 50 (200 I) . 55-62 0031-0174/2001/55-62 $200

The (): dendrology-, ecology and wood anatomy with reference to its applicability for tropical dendrochronology

M. TOMAZELLO F, P.C. BOTOSSO AND C.S. LISI

Department of Sciences, University of Slio Paulo, /34/8-900, Piracicaba, .

(Received 13 April 2000; revised version accepted 10 December 200 I)

ABSTRACT

Tomazello FM, Botosso PC & Lisi CS 2001. The genus Toona (Meliaceae): dendrology, ecology and wood anatomy with reference to its applicability for tropical dendrochronology. Palaeobotanist 50(1) : 55-62.

The potentiality of Too/w sp. trees, Meliaceae native in and continents, for dendrochronological studies is described, including its silviculture, ecology and wood anatomy. Emphasis is given to the presence of distinct annual growth rings sensitive to climatic variations and detectable by wood cross section analysis and X-ray densitometry technique.

Key-words-Toona sp .. Meliaceae, Dendrochronology. X-ray densitometry, Growth rings.

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INTRODUCTION 1976a; Styles & Khosla, 1976; Barroso, 1984). This family is represented by 8 genera in South America-, AXA under Meliaceae occurs in tropical and Swietenia, Cabralea, Trichilia, Guarea, Carapa, Schmardaea pantropical regions throughout Africa, America and and Ruegea - with the first 6 genera naturally occurring in T Asia, grouped in 51 genera and 1400 with Brazil. In addition exotic genera introduced from other regions approximately 500 species of economic importance are-Melia, Azadirachta, Toona and Khaya, etc. (Holdridge, (Lawrence, 1951; Styles, 1972; Girardi, 1975; Holdridge, 1976a; Rizzini,1978; Pennington, 1981; Barroso, 1984). The

© Birbal Sahni Institute of Palaeobotany, 56 THE PALAEOBOTANIST

Meliaceae family was studied by several researchers, like De varieties), T calanlas, Tfargesii, T lIlicrowrpa, T IllOllis, T Candolle (1878), Kribs (1930), Smith (1960, 1965), mullijuga, T paucijuga, T sarma, T serru!mQ, T sinensis Pennington and Styles (1975), Holdridge (1976a, b). In and T sureni. The most important species for the ecological addition there are several reprots from Brazil itself-Pirani conditions in Latin America are T ciliala val'. australis, T (1984), Pastore and Berzaghi (1989) in Sao Paulo, Amaral calanlas e T sareni. Actually, the better revision referring to (1981) in Goias, Klein (1984) in Santa Catarina, Girardi (1975) the genus Toona was done by Bahadhur (1988), in India, in Rio Grande do Sui and Pinheiro (1986) in Minas Gerais representing an analysis of the taxonomic knowledge, States. including descriptions, occurrence, economic uses, habitats, Among the representative species of Meliaceae, in this vernacular names, etc. paper emphasis is given to the genus Toona, considering its silvicultural and ecological importance, wood quality and Roem. var. australis (F. v M.) C. dendrochronological applications in tropical regions of the DC. world. Too/la ciliala has a natural widespread distribution, THE GENUS TOONA: DENDROLOGY, throughout India, Pakistan, Myanmar, Thailand. southern ECOLOGY AND WOOD ANATOMY , New Guinea. Malaysia, Philippines, Him31ayan valleys (up to 1300 m height). Molucas Island, etc. The trees The systematic classification of the genus Toona has can be found across the river banks 3nd in inclined lands. been discussed since long time by many authors, because of also in swampy 3nd wet tropic31 . including all the its similarity to the genus Cedrela. According to Smith western Pacific region (Grijpma & Ramalho. 1969). (1960),2 genera were initially established by Roemer in 1846: Tool1a cilia/a val'. aus/ralis trees naturally occurs in Cedre!a including the American cedars and Too/la, the eastern Australia in (UlladuI3, southern Asiatic cedars. Later, in 1878, De Candolle grouped them as Sidney) and States (nonhern Atherton) 3nd has Cedrela, because its close similarities and Harms, in 1896, the synonyms T alls/ralis, Cedrela aus/ralis and C. toona classified 2 different genera, regrouped again by De Candolle val'. australis. It's denominated "cedro australiano" or toona in 1908 (Pinheiro el ai., 1994) in Brazil, toona and cedar in Mexico, red cedar, Australian However classification of Harms was accepted by Smith toona, Australian red cedar in all spe3king English countries. (1960), pointing of the differences between Cedrela and The trees show monopodial growth and a superficial root Too/la. Chevalier and Begemann, cited by Grijpma and system, reaching 30-46 m in height and 1,50-2,00 m trunk Ramalho (1969) relates 11 species, Toona cilia/a (with 20 diameter and are light tolerant in the juvenile period,

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1 Fig. I-Curve of growth ri ng widlh of o ] , i t IIIIII II IiI Tool/a dlia[{l 1841 1851 1861 1871 1881 1891 1901 1911 1921 1931 19411951 1961 1971 1981 lreSS (afler Bhattacharyya year el al. 1992) TOMAZELLO elal-THEGENUS TOONA (MELIACEAE) DENDROLOGY. ECOLOGY AND WOOD ANATOMY 57

PLATE 1 Wood anatomy and growth rings of Tool1a cilia/{[ var. australis tree. Cross section of a trunk (a) and wood (b) showing growth rings. delimited by initial parenchyma bands and semi-porous ring. Cross section (c. d) showing the features of the parenchyma cells. vessel and fiber wall thickness in early and latewood. demanding sunlight in adult phase. In Australia it occurs in the ground water table is accessible to the roots in the dry low to median lands, wet to very wet climates. with 1l00­ period. The temperatures can vary fr,om -1 to 43°C, supporting 4000 mOlannual precipitation and a dry period of 3-4 months. light frosts; in Atherton region the temperature varies from -2 Despite their preference to wet soils. the trees can also develop to 28°C. Rich and well-drained soils are ideal for taona. which in driest regions. with 800 mOl annual precipitation, where prefers chalky soils and does not support dense clayish and 58 THE PALAEOBOTANIST poor sandy soils (Grijpma & Ramalho, 1969; Walters, 1974; fall in June-July (Pinheiro, 1986). Plantations cited by Rocas, 1986; Namikawa, 1988; Pinheiro et aI., 1994). Ledoux and Lobato (1976) in the Amazonian region prove In Central America countries, like Mexico the T ciliata the potential of toona. despite the great variability between trees were introduced from India and Myanmar and reach trees. In the same way, plantations in Sao Paulo and Espfnto /5-17 m height in tropical area plantations (Rocas, 1986). In Santo States demonstrate its potential concerning the wood Costa Rica the trees grow excellent in well-drained and volumetric increments (IPEF, 1975). The T ciliata val'. nutrient-rich soils with positive response to mineral australis descriptions, including , botany, ecological fertilization. The pruning of branches is recommended in aspects, silvicultural features, phenology. etc.. are presented 15-20 years old trees for higher wood quality (Otarola et al., by Gripjma and Ramalho (1969) and Pinheiro (1986). 1976; Sanchez et aI., 1976). The wood characteristics of T ciliata trees and its variety J Introduced in Hawaii in 1914, this tree species has been australis are similar, with specific gravity of 0'45-0'64 g!cm , cultivated for its high increment rates, reaching 30-36 m in presenting the same characteristics as Cedrela (Record & Hess. height and 0'25-0,65 m diameter in 22 years, producing wood 1947). Toona is considered as one of the best quality woods with excellent properties. Their phenology is characterized in India, Australia. etc., widely applied in furniture. carpentry. by the flowering in April-June, fruit ripening and seed cabinet, etc. The bark can be used in medicine and trees as dispersal in July-October and leaves shading in dry sites with ornamental in parks and gardens (Grijpama & Ramalho, 1969; hydric deficit (Walters, 1974). Walters, 1974; Rocas, 1986; Namikawa, 1988). In South America, Toona ciliata val'. australis trees were The wood of T ciliata presents sapwood rose-light brown introduced in 1969, in Argentina, by Cozzo and Mangieri, and in the val'. australis the color is yellowish-white, and the with aim to replace the cedar species in a reforestation heartwood darkened-brown to reddish. The wood anatomy program, however in Missiones Province, 10 year-old toona and properties were studied by many authors (Francis. 195 J; plantations suffered from photosynthetic problems (Mangieri, Bhat, 1985; Espinoza de Pernia, 1987; Cardoso & Tomazello 1972; Sanchez et aI., 1992). In 1988, 100na was introduced Filho, 1988; Sudo, 1989; Haslett er al., 1991), including fibers in the Alto Parana region, Paraguay, where the tree and early and latewood measurements (Bisset et al., J950). plantations show high growth increment rates compared to Additionally, dendrochronological studies were developed by other species (Serafina et aI., 1994). Chowdhury (1939). Bhattacharyya er at (J 992) and Cardoso In BraZil, toona shows good adaptation to several and Tomazello (J 988), due to the presence of distInct annual ecological conditions, also presenting resistance behaviour growth rings, delimitated by some typical anatomical wood against Meliaceae shoot borer, Hypsipyla grandella, while it features like initial parenchyma bands, vessel arrangement is attacked in its country of origin by H. robusta (Grijpma, and fiber wall thickness (PLATE J). 1976). In Minas Gerais State, toona trees reveal good growth increments and well-marked phenological events, flowering in September-November, fructification in January-March and

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Fig. 2--Chronology of growth ring width index of Toona ciliata trees (after Bhanacharyya et al., 1992). TOMAZELLO el 01.- THE GENUS TOONA (MELIACEAE): DENDROLOGY, ECOLOGY AND WOOD ANATOMY 59

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Fig 3-Chronology of growth ring width index and annual rain­ 1 I ' ~ ~t fall showing the correlation between Toolla ciliara tree 1 growth and annual rainfall (aft er Bhattacharyya el ai., 40 L t , I I I I I 1 """T , I 1(92) 1872 1882 1892 1902 1912 192219321942195210021972 1982 year

TOOlla sureni (BI.) Merril. Merril et Rolfe

This species naturally occurs in low lands up to 1000 m The species occurs in several provinces of the altitude in the dense pluvial equatorial forests of Vietnam, Philippines, near to the riverbanks and areas periodically Cambodia, Indonesia, Myanmar and Malaysia. The tree inundated. The trees reach 40-50 m height and up to 1·50 m shows high growth increments, reaching 20-40 m height and diameter. The straight and cylindrical trunk is free of 0,60-2,00 m diameter, producing high quality wood. The branches until 50% trunk height. Heartwood light to dark heartwood color is dark-brown, with characteristic odour, 0,39­ reddish, specific gravity 0'41-0'44 g/cm3 The wood call be 0'45 g/cm3 specific gravity, usually used in furniture, carpentry, easily worked and is applied in furniture, general carpentry, etc. The trees are strongly attacked by in musical instruments, etc. (Gripjma & Ramalho, 1969). Asia, when growing in plantations. However, the toona trees III Puerto Rico plantations are resistant against H. gnarled, TOONA CILIATA: A POTENTIAL SPECIES reaching 4·3 (2'30-8,30) m height and 7·0 (2'5-13-5) cm IN TROPICAL DENDROCHRONOLOGY diameter in 5·5 years (Gripjma & Ramalho, 1969). Toona ciliata val'. australis presents distinct annual growth rings constituted by fibers with length variations 60 THE PALAEOBOTANIST

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Fig. 4-Wood density profile by X-ray densitometry and demarcation of annual growth ring and boundaries of early (EW) and late wood (LW) of To(ma ciliata tree (ref., Text-Figure I). related to the diameter increment rates. The shorter fibers, at semi-porous ring, making possible the determination of tree the first earlywood layers, are formed in the beginning of age and annual increment rates (Cardoso & Tomazello Filho. spring, during the period of maximum cambium activity, while 1988). Besides direct growth ring measurements on the the latewood fibers of last layers produced in winter, during polished cross section, the X-ray densitometry method can be the minimum cambium activity are 83% longer (Bisset et al., applied for the delimitation of annual growth rings of toona 1950). wood samples. Significant values of maximum wood density In India, T ciliata is a deciduous tree, starting radial between and within growth rings also can be obtained, like 5° growth only after the renovation of 50% of the canopy with and 16° annual rings, with a specific gravity of 0·35 and 0·60 new leaves. In response, the cambium produces distinct g/cm3, respectively. These results show the potentiality of both annual growth rings, the boundaries of which are delimited by methodologies in the dendrochronological studies of tropical initial parenchyma bands and large diameter vessels forming species, i.e., Toona ciliata, including tree age determination, semi-porous rings, resulting in a period offast growth followed stand dynamics and relationship to climate, etc. (Fig. 4). by a period of lower growth rates (Chowdhury, 1940; Chowdhury & Rao, 1948). CONCLUSIONS Also in India, T ciliata trees are found potential for dendrochronological studies. They exhibit annual growth-rings Toona ciliata has a great potential for dendroclimatology sensitive to climatic variations. The growth rings were dated studies and forest dynamics in natural forests and plantations and measured and a reference-chronology of growth ring width in its area of natural occurrence and in Asia as well as in from 1800-1987 compared with growth ring width index tropical areas of Central and South America. In American chronology and annual rainfall. They recorded that the narrow countries, including Brazil, the phenological characteristics growth rings width was correlated with hydric deficit in the of introduced toona trees are similar to other native Meliaceae soil, caused by a low photosynthetic activity during the less species, mainly Cedrela and Swietenia. In these areas, rainfall years during the peak of monsoon rainy season (June­ however, resistant behaviour presented by toona trees against August), when growth is supposed to be fast. Toona trees the Meliaceae shoot borer, Hypsipyla grandella constitutes produce a superficial root system when growing in flat sticky the main advantage. In addition these species produces red soils with low water infiltration and deficient drainage, distinct annual tree rings delimitated by a typical initial when the water saturation may reduce soil oxygen supply and, parenchyma and semi-porous ring, essential for consequently, inhibit the root growth (Bhattacharya et al., dendrochronological studies. The wood anatomy and X-ray 1992) (Figs. I, 2, 3). densitometry methodologies can also be applied to the annual In Brazil, Toona ciliata trees produce distinct annual growth rings delimitation and to the intra- and inter-tree-ring growth rings characterized by initial parenchyma bands and density determination. However, a limited amount of TOMAZELLO <>( a{- THE GENUS TOONA (MELlACEAE): DENDROLOGY. ECOLOGY AND WOOD ANATOMY 61 dendrochronological work has been carried out on toona trees, Holdridge LR 1976a. Taxonomia de las meliaceas latino-americanas. In : Whitmore JL (Editor)-Studies on the schootborer II. despite of the confirmed sensitivity of the annual growth rings grandella Zeel. Turrialba. Costa Rica, CATIE 3 : 5-6. to climatic variation. Holdridge LR 1976b. Ecologia de las meliaceas latino-americanas ill : Whitmore JL (Editor)-Studies on the schootborer H. grandel/a Acknowledgements-The a/{Ihors are IhankJ/{1 10 Ihe FWldGi;ao Zeel. Turrialba. Costa Rica, CATIE 3 : 7p . de Amparo aPesquisa do ESlado de Sao Paulo-!'APESPJorflnallc­ IPEF - Instituto de Pesquisas e Estudos Florestais 1975. ing Ihe research projecl which originaled Ihis paper Desenvolvimento de programa de pesquisas: folhosas exoticas em regi6es tropicais e subtropicais. Boletim Informativo 3 : 45­ 51. REFERENCES Klein RM 1984. Flora ilustrada catarinense - meliaceas. Herbario "Barbosa Rodrigues". 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Serafina I, Barrienlos C & Jacquet B 1994. ESlUdio del Styles BT & Khosla PK 1976. Cytology and reproduclive biology comportamiento de la Toona ciliata en las condiciones del Alto of Meliaceae. In: Burley J& Tyles BT (Editors)- Tropical trces: Parana. Revista Forestal del Paraguay 10: 4-7. variation, breeding and conservation. London. Academic Press 2 Smith Jr CE 1960 A revision of Cedrela (Meliaceae). Fieldiana : 61-67. 29(5) : 295-341 Sudo S 1989. Wood anatomical characteristics of tropical species Smith Jr CE 1965. Meliaceae. In: Woodson RE (Editor)-Flora of from the Pacific region and Asia. In : Pacific Regional Wood Panama. Annals Missouri Botanical Garden 52 : 56-77. Anatomy Conference. Phillipines : 193·208. Styles BT 1972. The flower biology of the Meliaceae and its bearing Walters GA 1974. Toona australis Harms. In: Seeds of woody plants on tree breeding. Silvae Genetica 21 : 175-82. in United States. Forest Service. U.S. Department of Agriculture. Agriculture Handbook 450 : 813-814.