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Diametric Growth of Tree Species in the Atlantic Forest, Paraná, Brazil

Abstract

Abstract: Management of remnants in Atlantic Forest is an alternative for their conservation, however, information on the growth and ecology of those species is lacking. This study aimed to describe diametric growth of Balfourodendron riedelianum, Cordia trichotoma and Ocotea diospyrifolia based on its growth rings and to verify the relationship between this growth with the environmental characteristics in different altitude levels and forest types. Diametric growth was higher for the largest tree diameters of the three species. Based on the fitted growth model, the age in which mean annual increment in diameter becomes greater than the current annual increment was 55 years for B. riedelianum (DBH = 18.27 cm), 45 years for C. trichotoma (DBH = 26.56 cm) and 44 years for O. diospyrifolia (DBH = 26.05 cm). Environmental conditions and forest types affected diametric growth of these species. B. riedelianum and O. diospyrifolia showed higher diametric growth in plain regions with higher fertility soil and few frosts. C. trichotoma was negatively affected by low water availability in winter at lower altitudes and showed higher diametric growth at higher altitudes, where soil fertility is low but there is well-drainage, high rainfall and high relative humidity during the dry season.

Key words
Dendrochronology; altitude levels; forest types; forest management; forest ecology; Iguaçu National Park


INTRODUCTION

The Atlantic Forest is one of the most diverse biomes and is the home of many endemic species (Guedes-Bruni et al. 2009GUEDES-BRUNI RR, SILVA AG and MANTOVANI W. 2009. Rare canopy species in communities within the Atlantic Coastal Forest in Rio de Janeiro State, Brazil. Biodivers Conserv 18(2): 387-403.). The agricultural expansion followed by industrialization and urban development resulted in a significant degradation and fragmentation of this biome to small areas with less than 12.6% remaining of its original forest area (Ribeiro et al. 2009RIBEIRO MC, METZGER JP, MARTENSES AC, PONZONI FJ and HIROTA MM. 2009. The Brazilian Atlantic Forest: How much is left, and how is the remaining forest distributed? Implications for conservation. Biol Conserv 142(6): 1141-1153.).

In order to guarantee the conservation of the remnants, the Atlantic Forest Law was established and, among other rules, it prohibits the commercialization of native species. This rule has been established due to the lack of technical information on growth of native species and the technical and economic feasibility of small area management (BrasilBRASIL. 2006. Lei nº 11.428, de 22 de dezembro de 2006. Diário Oficial [da] República Federativa do Brasil. Poder Executivo, Brasília, DF. 26 dez. 2006. 2006). The lack of information on native species and their potential for economic use also contributed to the abandonment of forest remnants by the rural producers, as they consider these sites unproductive. In this context, efforts seeking to reveal how the growth of native species occurs are necessary for the Atlantic Forest.

Growth of forest species has been studied worldwide by the measurement of growth rings (MattosMATTOS PP, BRAZ EM, HESS AF and SALIS SM. 2011. A dendrocronologia e o manejo florestal sustentável em florestas tropicais. Colombo: Embrapa Florestas, Corumbá: Embrapa Pantanal, 37 p. et al. 2011, De RidderDE RIDDER M, BULCKE JV, ACKER JV and BEECKMAN H. 2013. Tree-ring analysis of an African long-lived pioneer species as a tool for sustainable forest management. For Ecol Manag 304: 417-426. et al. 2013, Vlam et al. 2014VLAM M, BAKER PJ, BUNYAVEJCHEWIN S and ZUIDEMA PA. 2014. Temperature and rainfall strongly drive temporal growth variation in Asian tropical forest trees. Oecologia, 174: 1449-1461., LatorracaLATORRACA JVF, SOUZA MT, SILVA LDSAB and RAMOS LMA. 2015. Dendrocronologia de árvores de Schizolobium parahyba (Vell.) S. F. Blake de ocorrência na rebio de Tinguá - RJ. Rev Árvore 39: 385-394. et al. 2015, BrandesBRANDES AFN, ALBUQUERQUE RP, MORAES LFD and BARROS CF. 2016. Annual tree rings in Piptadenia gonoacantha (Mart.) J. F. Macbr. in a restoration experiment in the Atlantic Forest: potential for dendroecological research. Acta Bot Bras 30(3): 383-388. et al. 2016, KanieskiKANIESKI MR, GALVÃO F, ROIG FA and BOTOSSO PC. 2017. Dendroecologia de Sebastiania commersoniana (Baill.) L. B. Sm. & Downs e Hovenia dulcis Thunb. em uma área degradada na floresta ombrófila mista aluvial, Sul do Brasil. Ciênc Florest 27(4): 1201-1215. et al. 2017). Climate-growth relationships are also extensively known (Costa et al. 2015COSTA MS, FERREIRA KEB, BOTOSSO PC and CALLADO CH. 2015. Growth analysis of five Leguminosae native tree species from a seasonal semidecidual lowland forest in Brazil. Dendrochronologia 36: 23-32., Rohner et al. 2016ROHNER B, WEBER P and THÜRIG E. 2016. Bridging tree rings and forest inventories: How climate effects on spruce and beech growth aggregate over time. For Ecol Manag 360: 159-169., Granato-Souza et al. 2018GRANATO-SOUZA D, ADENESKY-FILHO E, BARBOSA ACMC and ESEMANN-QUADROS K. 2018. Dendrochronological analyses and climatic signals of Alchornea triplinervia in subtropical forest of southern Brazil. Austral Ecol 43(4): 385-396.), contributing to identify the environmental preferences of a given species, for the restoration of degraded areas and to analyze the effects of climate changes.

The measurement of growth rings is an efficient technique that allows to obtain information on past diameter growth both quickly and at a low cost. For a growth ring reading to be reliable it must be performed annually or with known periodicity and distinguished by some anatomical feature of the wood (BotossoBOTOSSO PC and MATTOS PV. 2002. Conhecer a idade das árvores: importância e aplicação. Colombo: Embrapa Florestas, 25 p. and Mattos 2002); techniques such as cross-dating are still required to minimize errors in reading (Worbes 1995WORBES M. 1995. How to measure growth dynamics in tropical trees: a review. Iawa J, 16: 337-351.). Information about tree growth is used in forest management to determine the Biological Rotation Age (BRA) and the Minimum Logging Diameter (MLD) of each species (SchöngartSCHÖNGART J, WITTMANN F, WORBES M, PIEDADE MTF, KRAMBECK H and JUNK WJ. 2007. Management criteria for Ficus insipida Willd. (Moraceae) in Amazonian white-water floodplain forests defined by tree-ring analysis. Ann For Sci 64(6): 657-664. et al. 2007, 2008SCHÖNGART J. 2008. Growth-Oriented Logging (GOL): A new concept towards sustainable forest management in Central Amazonian várzea floodplains. For Ecol Manag, 252: 46-58., De Ridder et al. 2013, López et al. 2013). Toledo et al. (2011)TOLEDO M et al. 2011. Climate is a stronger driver of tree and forest growth rates than soil and disturbance. J Ecol 99: 254-264 also pointed out that knowing environmental conditions and your effect in tree growth rates contribute to forest management.

Balfourodendron riedelianum (known as Pau-Marfim), Cordia trichotoma (known as Louro-Pardo) and Ocotea diospyrifolia (known as Canela-Loura) are native species of the Atlantic Forest (LorenziLORENZI H. 1998. Árvores brasileiras: manual de identificação e cultivo de plantas arbóreas nativas do Brasil. 2ª ed., Nova Odessa: Plantarum, 352 p. 1998, CarvalhoCARVALHO PER. 2002. Louro-pardo - Cordia trichotoma. Circular Técnica 66. Colombo: Embrapa Florestas, 16 p. 2002, 2004CARVALHO PER. 2004. Pau-marfim - Balfourodendron riedelianum. Circular Técnica 93. Colombo: Embrapa Florestas, 11 p.). As these species have distinct rings formed annually (BoninsegnaBONINSEGNA JA et al. 1989. Studies on tree rings, growth rates and age-size relationships of tropical tree species in Misiones, Argentina. Iawa J 10(2): 161-169. et al. 1989, Maria 2002MARIA VRB. 2002. Estudo da periodicidade do crescimento, fenologia e relação com a atividade cambial de espécies arbóreas tropicais de florestas estacionais semideciduais. Dissertação (Mestrado) - Escola Superior de Agricultura “Luiz de Queiroz”. Universidade de São Paulo, 126 p. (Unpublished)., MattosMATTOS PP, TEIXEIRA LL, SEITZ RA, SALIS SM and BOTOSSO PC. 2003. Anatomia de madeiras do Pantanal mato-grossense: (características microscópicas). Colombo: Embrapa Florestas, 182 p. et al. 2003, Lisi et al. 2008LISI CS, TOMAZELO FILHO M, BOTOSSO PC, ROIG FA, MARIA VRB, FERREIRA-FEDELE L and VOIGT ARA. 2008. Tree-ring formation, radial increment periodicity, and phenology of tree species from a Seasonal Semi-deciduous Forest in Southeast Brazil. Iawa J 29(2): 189-207., Caum 2013), as well as a great value for the furniture industry, they were selected for this study, which consisted of the following objectives: (1) to describe growth patterns from growth rings diameter and; (2) to identify the effect of environmental characteristics (soil, temperature and rainfall) at different altitudes and forest types on the diametric increase.

MATERIALS AND METHODS

STUDY AREA

Iguaçu National Park (INP) is located in the western region of the state of Paraná, southern Brazil, and has a total area of 185,262 hectares (ha) (Figure 1). Köppen’s climate classification type is Cfa (AlvaresALVARES CA, STAPE JL, SENTELHAS PC, GONÇALVES JLM and SPAROKEV G. 2013. Köppen’s climate classification map for Brazil. Meteorol Z 22: 711-728. et al. 2013) with transition to Cfb in the northern region of the park. In the southern region, the historical average air temperature (1983-1997) ranges from 16°C (June) to 25.7°C (January) with an absolute minimum of -1.2°C (July), while the rainfall varies from 99.7 mm (July) to 227.6 mm (October). In the northern region, the historical average air temperature (1973-1998) ranges from 15.1°C (June) to 23.1°C (January) with an absolute minimum of -4.2°C (July), while the rainfall ranges from 108.7 mm (July) to 227.5 mm (October) (Iapar 2017IAPAR. 2017. Médias Históricas em Estações do IAPAR. Instituto Agronômico do Paraná. Disponível em: <http://www.iapar.br>. Acessado: 24 de Abril, 2017.
http://www.iapar.br>....
).

Figure 1
(a) Study area in the Brazil. (b) Iguaçu National Park in the Western of Paraná State.

The Iguaçu National Park is located in the Atlantic Forest Biome, with the Semideciduous Seasonal Forest (SSF) as the predominant vegetation (Souza et al. 2017SOUZA RF, MACHADO AS, GALVÃO F and FIGUEIREDO FILHO A. 2017. Fitossociologia da Vegetação Arbórea do Parque Nacional do Iguaçu. Ci Fl 27(3): 853-869.). In the southern region (<600 m), SSF Typical Submontane occurs in the plains, with deep soil of slow drainage and high fertility (Eutrophic Red Nitosol) (Souza et al. 2017). The SSF Humid Submontane occurs in this same altitude range, located on the slopes and base of the drainage ramps, with varying soil types (Eutrophic Red Nitosol, Eutrophic Melanic Tb Gleisol and Eutrophic Regolitic Neosol) and high humidity associated with the surrounding rivers (Souza et al. 2017).

In the northern region of the INP there are montane environments (≥600 m), with a predominance of well-drained soils with low fertility (Distrophic Red Latosol). The vegetation is classified as SSF Montane (600-700 m) and Ecotone between SSF Montane and Mixed Ombrophilous Forest (MOF) (>700 m) (Souza et al. 2017).

DATABASE

The field work was carried out at the end of the vegetative growth period in July 2015. Twenty-two trees of B. riedelianum with diameter at breast height (DBH) of 40.51±8.99 cm, 27 trees of C. trichotoma with DBH of 36.55±7.06 cm and 30 trees of O. diospyrifolia with DBH of 39.62±7.13 cm were sampled, with all measurements taken outside the bark. Around the permanent plots allocated in the study area described by Souza et al. (2017), the trees are distributed in the INP altitude and forest types according to Table I. The sampled trees were in the forest canopy, healthy and with tall and even stem. B. riedelianum presented height of the morphological inversion point of 13.94±2.98 m and total height of 19.51±4.17 m, C. trichotoma presented morphological inversion point of 14.29±2.92 m and total height of 20.00±4.09 m and O. diospyrifolia presented morphological inversion point of 10.67±2.83 m and total height of 14.94±3.96 m.

TABLE I
Trees sampled at altitude levels and forest types with respective environmentalcharacteristics in the Iguaçu National Park

In each sampled tree, two increment cores (90° angle) were collected with an increment borer (length 40 cm, diameter 5.15 mm) at DBH. Wood cores were glued on a wooden support, dried outdoors and polished consecutively with sandpaper from 100 to 600grit (Orvis and Grissino-Mayer 2002ORVIS KH and GRISSINO-MAYER HD. 2002. Standardizing the reporting of abrasive papers used to surface tree-ring samples. Tree-ring Res 58(1/2): 47-50.). Tree rings structure was analyzed by wood anatomical pattern using Leica DMS 300 stereomicroscopic (Figure 2). Maria (2002) and Lisi et al. (2008) delimited annual rings in B. riedelianum and described semiporous rings that differed by the combination of thick-walled fibers of late wood and marginal parenchyma, while emphasizing that fake rings are rare. Boninsegna et al. (1989) and Caum (2013) found rings formed annually in C. trichotoma, as well as semiporous rings distinct by late woods formed by small and medium sized vessels, thick-walled fibers and axial parenchyma. Finally, Mattos et al. (2003) described the formation of annual rings in O. diospyrifolia and rings marked by a darker tangential line with thick-walled radial fibers.

Figure 2
Wood anatomy of the species. Tree rings characterized in Balfourodendron riedelianum by the combination of latewood thick-walled fibers and marginal parenchyma (a) and (b); Cordia trichotoma by latewood made up of small and medium-sized vessels, thick-walled fibers and axial parenchyma (c) and (d); Ocotea diospyrifolia by a darker tangential line and radial thick-walled fibers (e) and (f). Arrows indicate the ring boundaries.

Wood cores were scanned with a HP Deskjet Ink Advantage 3636 at 600 dpi and ring widths were measured to the nearest 0.04 mm by ImageJ software (Nih 2016NIH. 2016. ImageJ version 1.50i. Disponível em: <https://imagej.nih.gov/ij>. Acessado: 16 de Setembro, 2016.
https://imagej.nih.gov/ij>....
) in the bark-pith direction. Cross-dating with COFECHA software (Holmes 1983HOLMES LR. 1983. Computer-assisted quality control in tree-ring dating and measurement. Tree-rings Res 43: 69-78.) and graphical analysis between wood cores of the same tree were used to minimize labeling error in the false rings.

DIAMETRIC GROWTH PATTERN

The combined width of the two rings observed from two cores collected on the tree represent the annual diametric increment; the DBH in each year was obtained by the addition of these values from tree pith. For trees in which it was not possible to identify the tree pith location, the annual increments were subtracted from the DBH measured in the field disregarding the bark thickness, until the last ring observed.

In order to describe in details the diametric growth pattern of each species, the increment values were classified by DBH-classes of 5.0 cm from the pith. A t-test was carried out to verify the effect of the size on the diametric growth by grouping the increments into two classes (DBH <20 and DBH ≥20 cm), as the largest classes had few measured rings.

Description of the species diametric growth using a well-defined model is useful for forest management and allow the estimation of the diametric growth, current annual increment in diameter (CAId) and mean annual increment in diameter (MAId) curves. These values are used to define the Biological Rotation Age (BRA), which is determined by using the basal area (LópezLÓPEZ JA. 2002. Árboles comunes Del Paraguay: ñande yvyra mata kuera. 2ª ed., Cuerpo de Paz: Asunción. 458p. et al. 2013) or volume (Schöngart et al. 2007, 2008, De Ridder et al. 2013). The Chapman-Richards sigmoidal growth model (Richards 1959RICHARDS FJ. 1959. A flexible growth function for empirical use. J Exp Bot 10(29): 290-301.), was adjusted for each species dataset. This model was selected for efficiency and flexibility to express the growth and production of biological variables (Zhao-gang and Feng-ri 2003ZHAO-GANG L and FENG-RI L. 2003. The generalized Chapman-Richards function and applications to tree and stand growth. J For Res 14(1): 19-26., CruzCRUZ JP, LEITE HG, SOARES CPB, CAMPOS JCC, SMIT L and NOGUEIRA GS. 2008. Curvas de crescimento e de índice de local para povoamentos de Tectona grandis em Tangará da Serra, Mato Grosso. Rev Árvore 32(4): 679-685. et al. 2008, Machado et al. 2015MACHADO AS, SOUZA RF, APARECIDO LMT, RIBEIRO A and CZELUSNIAK BH. 2015. Evolução das variáveis dendrométricas da bracatinga por classe de sítio. Cerne 21(2): 199-207., Figueiredo Filho et al. 2017FIGUEIREDO FILHO A, RETSLAFF FS, RETSLAF FS, LONGHI-SANTOS T and STEPKA TF. 2017. Crescimento e Idade de Espécies Nativas Regenerantes sob Plantio de Araucaria angustifolia no Paraná. Floresta Ambient 24: e00104814.). In the growth model, Y is the estimated diameter outside the bark (cm); I is the age (years); and b0, b1 and b2 are the estimated coefficients.

DBH and age data were used to fit growth models. Accuracy was verified by the coefficient of determination (r2), standard error (Sy) and distribution of residuals. The age of the trees in which the pith was not identified was estimated in 2015 by dividing DBH minus the bark by the mean diametric growth (WorbesWORBES M. 1995. How to measure growth dynamics in tropical trees: a review. Iawa J, 16: 337-351. 1995, Botosso and Mattos 2002, Stepka et al. 2014STEPKA TF, FIGUEIREDO FILHO A, MATTOS PP and MACHADO SA. 2014. Idade e dendrocronologia em árvores nativas de Araucária, Cedro e Imbuia no Sul do Brasil. In: Floresta com araucária - Pesquisas ecológicas de longa duração. 1ª ed., Curitiba: Multi-Graphics, p. 117-164.). The age of the trees with visible pith was estimated using this method and thus, a t-test was carried out to validate the procedure.

EFFECT OF ENVIRONMENTAL CONDITIONS

The effects of soil, temperature and rainfall on the diametric growth of the species were analyzed following the distribution of trees in the altitudes and forest types (Table I). Altitudes and forest types were defined as treatments in two completely randomized designs (CRD) and current increments in diameter were considered replications. The availability of light through canopy profiles affects the growth of trees in native forests (Hubbell et al. 1999HUBBELL SP, FOSTER RB, O’BRIEN ST, HARMS KE, CONDIT R, WECHSLER B, WRIGHT S and LAO SL. 1999. Light-gap disturbances, recruitment limitation, and tree diversity in a Neotropical Forest. Science 283: 554-557.) so that trees located in the understory have lower diametric growth (Hart et al. 2010HART JL, AUSTIN DA and VAN DE GEVEL SL. 2010. Radial growth responses of three co-occurring species to small canopy disturbances in a secondary hardwood forest on the Cumberland Plateau, Tennessee. Phys Geogr 31(3): 270-291.). In order to minimize the effect of lack of light in the first stages of growth, only the annual increments with DBH ≥20 cm were considered (minimum diameter of the forest canopy trees, which were considered subjectively) in the statistical analysis of the environmental effects on diametric growth (Vera et al. 2012VERA NE, SILVA F, CRISTÓBAL LL and GARCÍA D. 2012. Crecimiento de lãs Principales Especies de un Bosque Secundario de la Reserva de Guaraní, Misiones. Yvyrareta, 19: 14-22.). The nonparametric Kruskal-Wallis test was applied followed by multiple comparisons (Siegel and Castellan 1988SIEGEL S and CASTELLAN NJ. 1988. Nonparametric statistics for the behavioral sciences. Singapore: McGraw-Hill, 399 p.), since there is no normal distribution and homogeneity of variances (Bartlett’s test, P<0.05).

RESULTS

READING AND MEASURING GROWTH RINGS

The process for verifying error of labeling in the false rings using cross-dating and graphical analysis allowed rings synchronization between wood cores. The mean intercorrelation between core pairs was 0.39 (critical point 0.33, P>0.01) for B. riedelianum, with ages estimated from 54 to 138 years; 0.53 (critical point of 0.51, P >0.01) for C. trichotoma, with ages estimated from 27 to 78 years; 0.57 (critical point of 0.42, P>0.01) for O. diospyrifolia, with ages estimated from 29 to 79 years.

DIAMETRIC GROWTH PATTERN

Current annual increment in diameter for DBH-classes and the total of each species are presented in Table II. All three species presented growth rate increase with DBH increase and C. trichotoma had the maximum growth rate earlier than the other species (0.67 cm year-1, DBH-class 20-25 cm). The means of the three species for DBH-class ≥20 cm was higher than for the DBH-class <20 cm (t-test, P<0.01). A single tree of B. riedelianum showed DBH ≥50 cm (DBH = 72.57 cm) resulting in a CAId decrease for the highest DBH-class. The maximum CAId for B. riedelianum occurred in DBH-class of 30-35 cm (1.52 cm year-1), for C. trichotoma in DBH-class of 15-20 cm (1.59 cm year-1) and for O. diospyrifolia in DBH-class of 30-35cm (1.71 cm year-1).

TABLE II
Measured rings (n) and mean for current annual increment in diameter (CAId) grouped by DBH-class of 5 and 20 cm for three forest species in the Iguaçu National Park.

The tree pith was not identified in 24 out of 32 B. riedelianum trees, 25 out of 27 of C. trichotoma and 18 out of 30 of O. diospyrifolia, whose ages were estimated from the respective mean diametric increment (t-test, P=0.46). With these estimates and the values observed for the trees in which it was possible to identify the tree pith, growth models were fitted for each species (Table III).

TABLE
Fitted growth models for three forest species in the Iguaçu National Park.

Using these fitted growth models, a mean diametric growth curve, CAId and MAId curves were obtained for each species (Figure 3). The asymptotic yield of DBH is estimated by the coefficient b0 of the Chapman-Richards model (Richards 1959, Zhao-gang and Feng-ri 2003). The maximum CAId was estimated at ages 29 (0.36 cm year-1), 25 (0.63 cm year-1) and 24 years (0.64 cm year-1) for B. riedelianum, C. trichotoma and O. diospyrifolia, respectively. The maximum MAId occurs with 55 (0.33 cm year-1), 45 (0.59 cm year-1) and 44 (0.59 cm year-1) years for B. riedelianum, C. trichotoma and O. diospyrifolia, respectively. These latter ages represent the BRA of studied tree species as MAId was higher than CAId.

Figure 3
Individual diametric growth curves and mean curves fitted by growth models (black line), current annual increment in diameter (CAId) and mean annual increment in diameter (MAId) to Balfourodendron riedelianum, Cordia trichotoma and Ocotea diospyrifolia in the Iguaçu National Park.

EFFECT OF ENVIRONMENTAL CONDITIONS

Significant differences were observed for mean CAId values among altitude levels and forest types for the three species (Figure 4) (Kruskal-Wallis, P<0.05). The lowest altitudes are favorable to the current increment for B. riedelianum; the highest mean for current increment in diameter occurs at 250 m (0.50 cm year-1), which was statistically equal to 150 m (0.43 cm year-1). The lowest mean for B. riedelianum was observed between 650 and 350 m (0.38 to 0.41 cm year-1). Among the forest types, the highest mean CAId for B. riedelianum was observed in SSF Typical Submontane (0.42 cm year-1), which was similar to SSF Montane (0.38 cm year-1) (Kruskal-Wallis, P>0.05).

Figure 4
Means for current increment in diameter (DBH ≥20cm) of Balfourodendron riedelianum, Cordia trichotoma and Ocotea diospyrifolia at altitude levels and forest types in the Iguaçu National Park; SSF - Seasonal Semideciduous Forest, MOF - Mixed Ombrophilous Forest.

At high altitudes, higher growth of C. trichotoma was observed: 0.74 cm year-1 at 750 m against 0.62 cm year-1 at 150 m. The highest mean (0.74 cm year-1) among forest types was in Ecotone SSF/MOF, SSF Montane and SSF Typical Submontane, which was higher than in SSF Humid Submontane (0.52 cm year-1) (Kruskal-Wallis, P<0.05). In general, higher growth of O. diospyrifolia occurred between 150 and 550 m (0.73 against 0.61 cm year-1) (Kruskal-Wallis, P<0.05). The highest mean CAId was observed in SSF Typical Submontane (0.72 cm year-1), varying between 0.58 and 0.64 cm year-1 in the other forest types (Kruskal-Wallis, P>0.05).

DISCUSSION

Tree rings were used to describe the diametric growth pattern of B. riedelianum, C. trichotoma and O. diospyrifolia and to analyze the effect of environmental conditions on their diametric growth. Diametric growth was higher in the larger DBH-classes for the three species, as observed for other species in native forests (Clark and Clark 1999CLARK DB and CLARK DA. 1999. Assessing the growth of tropical rainforest trees: issues for forest modeling and management. Ecol Appl 9: 981-997., Silva et al. 2002). Hubbell et al. (1999) and Silva et al. (2002)SILVA RP, SANTOS J, TRIBUZY ES, CHAMBERS JQ, NAKAMURA S and HIGUCHI N. 2002. Diameter increment and growth patterns for individual tree growing in Central Amazon, Brazil. For Ecol Manag 166: 295-301. reported that trees with larger diameter occupying the forest canopy receive more light on the crown and have higher photosynthetic rates, causing increases in diametric growth.

Vera et al. (2012) found the same dependence between increment and size for adult trees of O. diospyrifolia in a secondary forest, describing higher periodic increments (five years) for larger trees (DBH ≥20 cm, 1.47 cm year-1) than for smaller trees (DBH <20 cm, 0.63 cm year-1). Mattos (2007)MATTOS RB. 2007. Produtividade e incremento de Cabralea canjerana (Vell.) Mart., Cedrella fissilis Vell. E Cordia trichotoma (Vell.) Arrab. Ex Steud., em floresta nativa no Rio Grande do Sul. Tese (Doutorado) - Pós-graduação em Engenharia Florestal. Universidade Federal de Santa Maria, 105 p. also emphasized a positive correlation between the periodic increment in basal area (cm² year-1) and DBH-classes for C. trichotoma, Cedrela fissilis and Cabralea canjerana, attributing the result to lower competition of the trees that occupy the forest canopy.

For a sustainable management of native forests, it is necessary to define a MLD for each species, based on the respective BRA (Schöngart 2008, De Ridder et al. 2013, López et al. 2013). Using fitted growth models and based on diameter, the BRA values were estimated in 55 years with MLD of 18.27 cm for B. riedelianum, 45 years with 26.56 cm for C. trichotoma and 44 years with 26.05 cm for O. diospyrifolia. López et al. (2013) determined higher biological rotation ages (≥80 years) and, consequently, higher DBH values (≥45 cm) for seven species in the Bolivian Savanna, which is justified by the use of the basal area (cm²) instead of the diameter. Schöngart (2008) also described biological rotation ages in volume greater than 60 years for 9 of 12 species studied (DBH’s ≥53 cm).

The lowest BRA values observed for B. riedelianum, C. trichotoma and O. diospyrifolia are associated with the use of diameter as a biological variable, commonly not used in the management of natural forests. However, the diameter is an alternative for the management of small forest remnants in South Brazil. In addition, the estimated DBH values for BRA were based on trees that grew without the application of silvicultural techniques, which might result in higher diametric growth rate, as observed by Venturoli et al. (2015)VENTUROLI F, FRANCO AC and FAGG CW. 2015. Tree diameter growth following silvicultural treatments in a semi-deciduous secondary forest in Central Brazil. Cerne 21(1): 117-123. for other semideciduous species. Bulfe (2008)BULFE NML. 2008. Dinâmica de clareiras originadas de exploração seletiva de uma floresta estacional semidecidual na provincia de Misiones, nordeste da Argentina. Dissertação (Mestrado) - Pós-graduação em Engenharia Florestal. Universidade Federal do Paraná, 84 p. (Unpublished). for example, described an increase of 0.25 cm year-1 in periodic increment after application of selective harvesting (6 years) for B. riedelianum, 0.19 cm year-1 for O. diospyrifolia and 0.15 cm year-1 for C. trichotoma as compared to the control (no treatment).

Observed results for forest stands also indicate that these species may have high diametric growth when trees are exposed to light from the early years. For B. riedelianum, Carvalho (2004) described MAId of 1.18 cm year-1 (7 years), Kubota et al. (2015)KUBOTA TYK, MORAES MA, SILVA ECB, PUPIN S, AGUIAR AV, MORAES MLT, FREITAS MLM, SATO AS, MACHADO JAR and SEBBENN AM. 2015. Variabilidade genética para caracteres silviculturais em progênies de polinização aberta de Balfourodendron riedelianum (Engler). Sci For 43: 407-415. reported MAId between 0.55 and 0.59 cm year-1 (27 years) in the progeny test, and MattosMATTOS PP, BOTOSSO PC, CORDELLI RL and CARVALHO PER. 2004. Estudo dos anéis de crescimento de diferentes procedências de Balfourodendron riedelianum, Rutaceae e Cordia trichotoma, Boraginaceae, sob condições de plantio. In. 55º Congresso Nacional de Botânica e 26º Encontro Regional de Botânicos de MG, BA e ES. Viçosa. Livro de Resumos...Viçosa: CNBOT, p. 845. et al. (2004) found MAId of 1.0 and 1.15 cm year-1 for 12 and 15 years, respectively. For C. trichotoma, Radomski et al. (2012)RADOMSKI MI, PORFÍRIO-DA-SILVA V and CARDOSO DJ. 2012. Crescimento de Louro-pardo (Cordia trichotoma) Vell. Arráb. Ex Steud. em sistema agrossilvipastoril. In. VII Congresso Latinoamericano de Sistemas Agroflorestais para a Produção Pecuária Sustentável - Centro Brasileiro de Pecuária Sustentável. Belém do Pará, Anais...Belém do Pará: CBPS, p. 492-496. and Carvalho (2002) reported MAId values of 1.26 cm year-1 (7 years) and 3.07 cm year-1 (3 years), respectively.

Environmental characteristics such as soil, water availability, temperature and rainfall affect the distribution of forest species (Oliveira-Filho and Fontes 2000OLIVEIRA-FILHO AT and FONTES MA. 2000. Patterns of Floristic Differentiation among Atlantic Forest in Southeastern Brazil and the Influence of Climate. Biotropica 32: 793-810., BotrelBOTREL RT, OLIVEIRA FILHO AT, RODRIGUES LA and CURI N. 2002. Influência do solo e topografia sobre as variações da composição florística e estrutura da comunidade arbórea-arbustiva de uma floresta estacional semidecidual em Ingaí, MG. Braz J Bot 25(2): 195-213. et al. 2002, Ferreira-Júnior et al. 2007FERREIRA-JÚNIOR WG, SILVA AF, SCHAEFER CEGR, MEIRA NETO JAA, DIAS AS, IGNACIO M and MEDEIROS MCMP. 2007. Influence of Soils and Topographic Gradients on Tree Species Distribution in a Brazilian Atlantic Tropical Semideciduous Forest. Edinb J Bot 64(2): 137-157.), as well as the growth pattern (Toledo et al. 2011). With the reduction of altitude in the INP, where the winters are dry, with few days of rain and frost of low intensity, and the summers are hot and humid (Iapar 2017), B. riedelianum and O. diospyrifolia grew up better (Table II). At low altitudes, the highest diametric growth of both species occurred in SSF Typical Submontane, in plains with deep soil of slow drainage and high fertility (Eutrophic Red Nitosol) and lower humidity when compared to the SSF Humid Submontane (Souza et al. 2017).

The environmental conditions that favored the growth of B. riedelianum in the INP were also described by Carvalho (2004), corroborating the results of the present study. For O. diospyrifolia, this is the first study about the effect of environmental conditions on its diametric growth. It is noteworthy that this species showed higher growth rate between 150 and 550 m, with the exception of 350 m. This can be explained by the high humidity and shallow soil from the site at 350 m, which is characterized by SSF Humid Submontane, located in the slope of the drainage ramp and close to a river. Despite this result, due to the wide geographic distribution of O. diospyrifolia, occurring from north to south of Brazil (QuinetQUINET A, BAITELLO JB, MORAES PLR, ASSIS L and ALVES FM. 2015. Lauraceae na Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. Disponível em: <http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB8499>. Acessado: 14 de Junho, 2018.
http://floradobrasil.jbrj.gov.br/jabot/f...
2015), Paraguay (LópezLÓPEZ L, VILLALBA R and BRAVO F. 2013. Cumulative diameter growth and biological rotation age for seven tree species in the Cerrado biogeographical province of Bolivia. For Ecol Manag 292: 49-55. 2002) and Argentina (Vera et al. 2012), further investigations are necessary to corroborate the results found in this study.

In fact, the environmental influence on growth rates of forest species must be carefully analyzed. For instance, C. trichotoma presented higher diametric growth at the highest altitude in Ecotone SSF/MOF (Figure 3) with well-drainage soil of low fertility (Distrophic Red Latosol), while Carvalho (2002) reported that this species grows up in sites with well-drainage higher fertility soil. Caum (2013) reported a significant reduction on the CAId associated to the water deficit for this species, which explains, along with the well-drainage soil, the higher diameter growth at high INP altitudes, where there is higher rainfall and relative humidity during the dry season (Iapar 2017).

CONCLUSION

In summary, this study contributes to the management and conservation of the Atlantic Forest, providing ecological information and diametric growth for Balfourodendron riedelianum, Cordia trichotoma and Ocotea diospyrifolia, native species of commercial importance. Fitted models provided the Biological Rotation Age based on the diameter and respective Minimum Diameter Logging for each species in the Iguaçu National Park (55 years with 18.27 cm for B. riedelianum, 45 years with 26.56 cm for C. trichotoma and 44 years with 26.05 cm for O. diospyrifolia). The environmental conditions of SSF Typical Submontane (<600 m) favored the diametric growth of B. riedelianum and O. diospyrifolia. On the other hand, C. trichotoma presented higher diametric growth in the Ecotone SSF/MOF (>600 m), where trees face well-drained soils with low fertility, high rainfall and high relative humidity during the dry season.

ACKNOWLEGMENTS

We would like to thank the Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) for the authorization and availability of the physical structure to carry out this study. We would also like to thank the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the financial support in the form of scholarship.

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Publication Dates

  • Publication in this collection
    11 Nov 2019
  • Date of issue
    2019

History

  • Received
    21 Feb 2018
  • Accepted
    2 Apr 2019
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