IAWA Bulletin n.s., Vol. 10 (2),1989: 161-169

STUDIES ON RINGS, GROWfH RATES AND AGE-SIZE RELATIONSHIPS OF TROPICAL TREE SPECIES IN MISIONES,

by

J.A. Boninsegna*, R. Villalba*, L. Amarilla**, and J.Ocampo**

Summary Wood samples of 13 tree species from Several studies have been carried out on three sites in the Selva Misionera (Misiones the wood anatomy of tropical in order to Province, Argentina) were analysed macro­ identify the growth layers and their temporal and microscopically for occurrence and for­ sequence and several methods have been mation of growth rings. Well-defined annual used to reach this objective (Mariaux 1967; tree rings were found in Cedrelafissilis VeIl., Catinot 1970; Tomlinson & Craighead 1972; rigida Benth., Cordia tricho­ Eckstein et al. 1981; Lieberman et al. 1985; toma VeIl. and Chorisia speciosa St. Hil. Villalba 1985; Worbes 1985, 1986). In 15 trees of Cedrela fissilis VeIl., the Growth rates of woody as indicated growth rate, the current and the mean annual by their annual ring widths are always to increment (CAl and MAl) and the diameter/ some degree a function of both natural and age relationship were estimated using incre­ anthropogenic conditions (Fritts 1976). Com­ ment borer samples. The estimated mean cul­ parisons of ring widths in the same species mination age of the basal MIA was 152 over time or in different places can provide years, while the same parameter measured on valuable information on how woody individual trees shows a wide range from 61 growth varies temporally or spatially as a to 180 years, probably representing different function of various environmental conditions. social positions of the trees. In addition, it is possible to evaluate the re­ The method proposed is discussed as a lative productivity of species and sites, or way to obtain a quick, reliable, and inexpen­ how a species' growth is affected by various sive estimation of parameters which are use­ management practices. ful in forest management. The Selva Misionera represents the south­ Key words: Tropical trees, growth rate, tree ernmost part of the Amazon pluvial forest, rings. extending over Misiones Province, Argen­ tina, from 25° to 27° L.S. (Fig. 1). Precipi­ Introduction tation is distributed throughout the year One of the most frequent uses of tree ring reaching a total average of more than 1700 information is in the determination of the age mm. Mean annual temperature ranges be­ of individual trees or stands of trees. Age de­ tween 19 to 20°C. Frosts occasionally occur, termination requires that a researcher is able mainly in June or July. Lateritic soils, in to recognise rings and also that the trees combination with high temperature and pre­ regularly produce one and only one ring each cipitation allow the growth of subtropical year. When a species is shown to have iden­ evergreen forest, called Selvas Mixtas (Ca­ tifiable annual rings, a ring count can nor­ brera 1976). The most important families are mally provide age estimates that are suffi­ Leguminosae, Lauraceae, Meliaceae, Boragi­ ciently accurate for many purposes (Duever naceae and Bignoniaceae (Takao Inoue et al. & McCollom 1987). 1984).

* Laboratorio de Dendrocronologfa, Centro Regional de Investigaciones Cientfficas y T6cno­ l6gicas, CONICEf, Casilla de Correo 330, 5500 Mendoza, Argentina. ** Facultad de Ciencias Forestales, Universidad de Nacional de Misiones, 3380 Eldorado, Misiones, Argentina.

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~ Sample site .uJJ Mesopotamico park· like forest C=:J Su btropical evergreen forest of Brasil _ Parana fluvial forest

_ Araucaria angustifolia forest I " d Pampas grassland

Fig. 1. Location map of northeastern Argentina and surrounding areas showing the sample site positions.

In this report, 13 species of the Selva 1. Ring-porous or semi-ring-porous struc­ Misionera are analysed in order to determine ture: their dendrochronological characteristics. On a. The earlywood consists of large ves­ one of them, Cedrela fissilis VeIl., a prelim­ selsembedded in a mass of initial pa­ inary study of the age and growth rate was renchyma cells: undertaken on the basis of its growth ring Cedrelafissilis (Plate la-c) widths. b. The latewood is made up of small and medium-sized vessels and thick- walled fibres: Cordia triclwtoma (Plate 2a, b) Wood anatomy and tree rings 2. Boundaries with uni- or multi seriate mar­ Anatomical studies of wood samples of 13 ginal parenchyma bands. The parenchyma tree species from different places in Misiones cells at the boundaries are often filled with Province were carried out with special atten­ crystalline substances: tion to the growth zones. The main anatomi­ Parapiptadenia rigida (Plate Id-t) cal structures responsible for the visibility of Peltoplwrum dubium (Plate 2d) growth rings are: Myrocarpus frondosus (Plate 2e, t)

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Plate 1. Woods with annual growth rings. - a-c: Cedrelafissilis. a: Transverse section show­ ing growth ring boundaries marked by continuous paratracheal parenchyma bands in combina­ tion with semi-ring-porosity. - b: Enlargement of the initial parenchyma band. - c: Details of the growth ring boundary showing thick-walled fibres in the latewood (lower part) and thin-walled parenchyma cells in the earlywood (upper part). - d-f: Parapiptadenia rigida. d: Cross section showing a marginal parenchyma band filled with amorphous substances at the growth ring boundary. - e: Details of the parenchyma band without cell contents. - f: Transverse section of secondary xylem with three complete zones. Annual rings are indicated by arrows.

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Plate 2. Woods with annual growth rings (continued). - a, b: Cordia trichotomfl. a: Tree rings defined by a narrow initial parenchyma band. In b, the parenchyma band is associated with semi-ring-porosity (arrows). - c: Chorisia speciosa showing an annual ring consisting almost solely of fibres in the earlywood. - d-f: Species with growth rings, not necessarily aTlnual. - d: Peitophorum dubium with irregular zonate bands of apotracheal parenchyma (arrows). - e, f: Myrocarpusfrondosus. e: Narrow parenchyma bands completely surrounding the trunk.­ f: Details of the 1-4-seriate parenchyma band.

---+ Plate 3. Species with less distinct, ± irregular growth increments. - a: Aspidosperma poiyneu­ ron, annual ring defined by a narrow band of fibres representing the first formed earlywood. - Nectandra sa/igna (b) and Ocotea puberuia (c) showing both gradual and abrupt changes in radial diameter and wall thickness of the fibres. - d-f: Enteroiobium contortisiliquum.

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d: Two interannual parenchyma bands (indicated by arrows) formed during the same year of growth. Growth rings defined by a reduction in radial diameter of the last formed fibres (e) or by a parenchyma band (t). - Species with indistinct growth rings, g: Balfourodendron riedelia­ num, and h: Cabralea oblongifolia.

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3. Growth zone boundaries conslsttng of The patterns described here may occur in several rows of fibre cells with a short various combinations within the growth layer radial diameter and thick walls. of some species. Thus, thin bands of paren­ a. Zones of thicker-walled fibres are pres­ chyma beside flattened fibre cells occur in the ent at the beginning of growth layers: same growth layers of Enterolobium contorti­ Chorisia speciosa (Plate 2c) siliquum (plate 3d-f). Aspidosperma polyneuron (plate 3a) The abrupt change from the late- to the Dendrochronological studies on the end earlywood with relatively low and high grains of disks and increment borer samples fibre percentages respectively, marks of the same material analysed microscopi­ the tree ring limit. Moreover, the tissue cally, have allowed to establish a prelimi­ percentage taken up by fibres gradually nary classification in relation to visibility of diminishes along the annual ring. growth layers, continuity along the cross sec­ b. Thicker fibre zones are found at the tion, and annual nature of growth zones. The end of growth layers. In this case, the annual nature was established taking into ac­ transition of growth zones can be grad­ count the studies performed on other species ual or abrupt in the same tree: of the same genus (Villalba et al. 1985) and Ocotea puberula (plate 3b) the local rate of tree growth according to Ma­ Nectandra saligna (plate 3c) radei (1982), Maradei et al. (1982), Gartland 4. Periodical recurring patterns of parenchy­ et al. (1969), Fernandez Rodriguez (1963), ma along the growth layers: and Mangieri (1957). These observations Cabralea oblongifolia (plate 3h) have been summarised in Table 1.

Table 1. Dendrochronological characteristics of some tropical trees growing in Misiones Province, Argentina.

Growth layers well defined Continuous over the entire cross section Representing one year of growth (plates 1a-f, 2a-c) Cedrelafissilis Vell. (Meliaceae) Parapiptadenia rigida Benth. (Legum.-) Cordia trichotoma Vell. (Boraginaceae) Chorisia speciosa St. Hil. (Bombacaceae) Not representing annual growth (Plate 2d-f) Peltophorum dubium Spreng. (Legum.-Caesalpinioideae) Myrocarpus frondosus Fr. Allem. (Legum. -Papilionoideae) Not continuous over the entire cross section Representing one year of growth (Plate 3a) Aspidosperma polyneuron Mull. (Apocynaceae) Sometimes, but not always representing one year of growth (Plate 3b-f) Ocotea puberu/a Nees. et Mart. (Lauraceae) Nectandra sa/igna Nees et Mart. (Lauraceae) Enterolobium contortisiliquum VeIl. (Legum. -Mimosoideae) Growth layers vague or absent (plate 3g, h) Tabebuia ipe Mart. (Bignoniaceae) Balfourodendron riedelianum Eng!. (Rutaceae) Cabralea oblongifolia C. DC. (Meliaceae)

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em 40 a

em b 80

60

40

20

30 60 90 120 150 180 210 50 100 150 200 age (years) age (years) em m2 40 0.4 c d

30 60 90 120 150 30 60 90 120 150 em age (years) age (years) 30 e f 40

30 20 MAl MAl CAl 20

10 10 CAl Ot:;.J...J~L....L..JL....L..J....LJ....LJ...J....1...J....1 30 60 90 120 150 30 60 90 120 150 age (years) age (years)

Fig. 2. Growth rate of Cedrela fissilis. a: Cumulative radial increment of individuals, mean curve = broken line. b: Age-diameter relationship. c & d: Mean cumulative radial and basal increment Mean annual increment (MIA) and current annual increment (CAl) from radial (e) and basal (f) data, respectively.

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Age and growth rate of CedreIa fissilis trees have not reached the culmination age A total of 15 trees of CedreLafissilis Vell. yet. This result could indicate differences in were cored twice at breast height Cored sam­ social position between the trees in the forest. ples were processed following Stokes and A more or less linear relationship could be Smiley (1968). The rings were observed established between age and diameter, with a under a microscope and the estimated age high correlation coefficient (r = 0.87). These of each one was recorded, starting from the results are of a preliminary nature because pith which was assigned the age of 1. Ring they are based on a very limited number of width measurements were obtained for every trees. growth year to the nearest one hundredth of a millimetre. The arithmetic mean of the values for each Conclusions ring was obtained and the series was smooth­ Several tropical trees are capable of form­ ed by a cubic spline filter of 0.65 N pass, in ing growth rings. Out of 13 species, macro­ which N indicates the number of elements of and microscopically analysed, from the Selva the series. Misionera, 10 species show well-defined Based on the ring width' of the corre­ growth rings, whereas only 3 species have sponding age and the previous radius, basal vague growth rings or lack them. The prin­ area increment (cm2) was estimated for each cipal anatomical structures involved in their growth year according to the equation: formation are initial parenchyma cells, termi­ nal parenchyma cells and radially flattened Ba (t) = 21t r (t-1) Ir (t) and/or thick-walled fibres at the beginning or in which Ba(t) is the basal area at age t, r(t-l) at the end of tree rings. The anatomical struc­ is the radius reached by the tree at age t-l ture responsible for the visibility of growth and Ir (t) is the ring width of the t-aged ring. rings is similar within species of the same Data analysis was twofold, involving the genus. CedreLafissilis from the Selva Misio­ comparison of radial and basal area growth nera as well as C. angustifolia Sesse Moc. and the determination of the following para­ and C. Lilloi C. DC., growing in the Tucu­ meters: mean annual radial, mean annual ba­ mano-oranense forest, show tree rings de­ sal, current annual radial and current annual fined by an initial band of parenchyma basal increment. The mean annual increment (Villalba et aL. 1985). This rule is not valid (MAl) is defined as the average annual in­ for species of the same family. Even though crement that occurs up to a given age as CabraLea obLongifolia belongs to the same calculated by dividing the accumulated in­ family as CedreLa fissilis (Meliaceae), the crement by the age. The current annual incre­ growth rings in the former species are vague ment (CAl) is the amount of growth that or not well defined. However, the occurrence occurs at a specified age. The culmination age of certain ring types is predominant in some is the age at which the MAl reaches its maxi­ families. The Leguminosae species analysed mum value (Assman 1970). mainly have bands of terminal parenchyma, The results are summarised in Figure 2a-f. even when in some species these bands Cross dating was not carried out on this are not necessarily annual. In Leguminosae material due to the low circular uniformity species in Central Amazonia, Worbes (1985) observed in cross section and the fact that it found that marginal parenchyma bands are was not needed in the analysis of the data. the most common anatomical structure in The differences in culmination age for ra­ relation to growth ring demarcation. dial increment (c. 63 years) and basal area The dendrochronological studies on Ce­ increment (c. 153 years), observed in the drela fissilis enable a test of the applicability mean curves, are particularly striking. of the tree ring width measurements in deter­ The analysis of the culmination age of the mining age and growth rate in a tropical tree. basal MAl made on individual trees, showed One important advantage of the method is that a wide variation, ranging from 61 to 180 it permits an insight into the dynamics of years, independent of tree diameter. Some growth of each individual and to a certain

Downloaded from Brill.com09/29/2021 08:25:25PM via free access Boninsegna, Villalba, Amarilla & Ocampo - Tree rings in species in Misiones, Argentina 169 degree into the life history of the tree. Al­ Hueck, K. 1978. Los bosques de Sudame­ though the age values have some errors due rica. Soc. Alemana de Coopemci6n. Tec­ to the different time at which individual tree­ nica, Eschborn. lets reach breast height, the determination of Lieberman, D., M. Lieberman, G. Hartshorn age and growth rate on the basis of tree ring & R. Peralta. 1985. Growth rate and age­ widths is a very economic, fast and reliable relationships of tropical wet forest trees in method that could be applied in tropical Costa Rica. J. Trop. Ecol. 1: 97-109. regions where there are no other sources of Mangieri, H.R. 1957. Notas sobre carac­ information. terfsticas y cultivo del Cedro misionero (Cedrela tubifiora). Rev. For. Arg. 1 (3): 112-114. References Maradei, D. 1982. Cultivo de especies de la Assman, E. 1970. The principles of forest Selva Misionera I. Resefia Bibliografica. yield study (ed. P.W. Davis). Pergamon Actas Primeras Jornadas Tecnicas sobre Press, New York. Bosques Implantados en el Noreste Ar­ Cabrera, A. C. 1976. Regiones Fitogeografi­ gentino, Un. Nac. de Misiones: 105-111. cas . In: Encidopedia Argentina - , A. Morales, J.R. Ruiz & J.E. Torres. de Agricultura y Jardinerfa 2. Edit. ACME, 1982. Ibid. II. Comportamiento inicial Buenos Aires. bajo cubierta de pinos. Ibid.: 111-113. Catinot, R. 1970. Premiers reflexions sur Mariaux, A. 1967. Les cernes dans les bois une possibilite d'explication physiologi­ tropicaux africains, nature et periodicite que des rhytmes annuels d'accroissement I: Peuvent-ils reveler l'age des arbres? II: chez les arbres de la foret tropicale afri­ Periodicite des cernes, methodes d' etude et caine. Bois et Forets Tropiques 131: 3- premiers resultats. Bois et Forets des Tro­ 14. piques No. 113: 3-14 & No. 114: 23-37. Duever, M. & 1. McCollom. 1987. Trade Stokes, M.A. & T.L. Smiley. 1968. An in­ offs between the use of tree ring counting troduction to tree ring dating. Univ. Chi­ and dendrochronology in ecological stud­ cago Press, Chicago. ies. Proc. International Symp. Ecological Takao Inoue, M., C. V. Roderjan & Y. S. Aspects of Tree Ring Analysis, Mary­ Kuniyoshi. 1984. Projeto Madeira de mount College, Tarrytown, New York: Parana. Curitiba, Fundaya5 de Pesquisas 611-621. Horestais do Parana. Eckstein, D., J. Odgen, G.C. Jacoby & J. Torillinson, P.B. & F.C. Craighead. 1972. Ash. 1981. Age and growth rate determi­ Growth-ring studies on the native trees of nation in tropical trees: The application subtropical Horida. In: Research trends in of dendrochronological methods. In: Age plant anatomy (eds. A.K.M. Ghouse & and growth rate of tropical trees: new di­ M. Yunus): 39-51. McGraw Hill, New rections for research (eds. F.H. Bormann Delhi. & G. Berlyn): 83-106. Yale Univ., New Villalba, R. 1985. Xylem structure and cam­ Haven. bial activity in Prosopis flexuosa. IAWA Fernandez Rodriguez, M. 1963. Multiplica­ Bull. n.s. 6: 119-130. ci6n espontanea del Peteribi (Cordia tri­ - ,1. Boninsegna & R. Holmes. 1985. Ce­ chotoma) previa eliminaci6n del bosque drela angustifolia and Juglans australis: original en la provincia argentina de Misi­ two new tropical species useful in dendro­ ones. Rev. For. Arg. 7 (4): 11-14. chronology. Tree Ring Bull. 45: 25-35. Fritts, H.C. 1976. Tree rings and climate. Worbes, M. 1985. Structural and other adapta­ Acad. Press, New York. tions to long-term flooding by trees in Cen­ Gartland, H.M. & C.M. Volkart. 1969. De­ tral Amazonia. Amazoniana 9: 459-484. terminaci6n del crecimiento de una planta­ - 1986. Lebensbedingungen undHolzwachs­ ci6n de Cordia tricotoma en la provincia tum in zentralarnazonischen Uberschwem­ de Misiones, Argentina. Actas 1er. Con­ mungswaldern. Scripta Geobotanica 17: greso Forestal Argentino: 223-225. 1-112.

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