Litterfall and Litter Decomposition in Pinus and Native Forests

ABSTRACT

Litter production and decomposition makes it possible to supply a good part of the nutrient demand of forest stands. Although several studies on this subject have been carried out in Pinus stands in different regions of Brazil, there are no records of studies carried out in the Northeast region, or in particular in the state of Bahia. Therefore, the objective of this work was to evaluate the litter production (litterfall) and decomposition in Pinus stands in the southwest region of Bahia, using an area of native forest as reference. Litterfall was evaluated during twelve months with the use of suspended collectors. Litter accumulation was quantified by means of three collections. The values obtained from litterfall were similar among Pinus stands. However, these values were lower than those found in the reference forest and in stands of the same genus in other regions of Brazil. Litter decomposition is relatively slow in Pinus sp. stands. The temporal variation of the litter supply is not very sensitive to the climate variations.

Keywords:  Reforestation; deposition; production; senescent material

1. INTRODUCTION

The forestry sector in Brazil has grown in the last decades. The progressive demand for products and by-products of forest origin such as oils, essences, and especially wood have provided a large increase in the number of reforestation areas with exotic fast-growing species (ABRAF, 2013). A typical example of this trend is the Pinus genus, the main source of raw material for several industries in the country, which has its most extensive plantations established in the states of Paraná and Santa Catarina, although it is also distributed in several other regions of the country, including the Northeast region (IBÁ, 2015).

According to the Brazilian Agricultural Research Corporation (Empresa Brasileira de Pesquisa Agropecuária – EMBRAPA, 2014), the Pinus genus comprises more than 100 species with great potential for several purposes. Its wood is mainly used in sawing and laminating, sheet metal, and pulp and paper industries. The main characteristics that have led to the wide use of this genus are wood quality, rusticity and tolerance, which enable its planting in soils which are marginal to agriculture (Chaves & Corrêa, 2005). In addition to the use of wood for various purposes, Pinus species excel in the production of resin used in glue, varnish, paint and adhesive industries (Medeiros et al., 2017).

Native or planted forests such as Pinus plantations present a continuous supply of senescent material from its aerial parts and root system, which favors the formation of an organic layer on the soil surface, also known as litter. According to Vieira & Shumacher (2010a), plant litter consists of vegetable debris (leaves, branches, stems, bark, fruit and flowers) and animal debris. This layer plays an important role in maintaining or increasing the production of forest ecosystems, since it allows a return of nutrients and carbon from the vegetal biomass into the soil through decomposition (Giácomo et al., 2012; Godinho et al., 2013; Pinto et al., 2016). It also plays an important role in the interception and retention of rainwater, thus contributing to an increase in the infiltration rate and water storage in the soil (Mateus et al., 2013). Therefore, plant litter together with the aerial and root parts of plants provide protection to the soil and can provide conditions for reestablishing its physical, chemical and biological properties (Andrade et al., 2000).

The presence of litter on the surface of forest soils and its consequent reuse in nutrient cycling, along with the decomposition process, enables the supply of a large part of the trees’ nutrient demands, thus guaranteeing an ecosystem functioning. The importance of this process is evidenced in forests found growing in soils of low fertility (Schumacher et al., 2003), which is the case of the great majority of areas planted with fast growing exotic forest species such as the Pinus genus in Brazil.

Although nutrient cycling is a common process for all forest ecosystems, the amount of litter produced and its decomposition rate are particular to each ecosystem. This pattern is determined by vegetation characteristics such as species composition and the quality of plant residue, in addition to the action of climatic variables which may alter the amount of litter deposited and advance (or not) its decomposition (Vieira & Shumacher, 2010b; Barbosa et al., 2017).

Several studies on litterfall and litter decomposition of Pinus stands have been carried out in different regions of Brazil (Carvalho et al., 2008; Schumacher et al., 2008; Vieira & Shumacher, 2010a; Piovesan et al., 2012; Lima et al., 2015). However, there are no records in the literature of studies carried out in the Northeast region, or in particular in the state of Bahia.

In view of the above, the present study had the objective to evaluate the litterfall and litter decomposition in two Pinus stands; a pure stand (Pinus caribaea), and another among a consortium (Pinus oocarpa, Pinus caribaea and Pinus taeda), both in the Southwest region of Bahia, as well as considering the possible interference of climatic variables. A semi-deciduous seasonal forest was used as a reference. Therefore, it was assumed that the dynamics of litterfall (production) and litter decomposition would differ according to the organization of the Pinus sp. stands (homogeneous or heterogeneous) and their climatic conditions.

2. MATERIAL AND METHODS

2.1. Characterization of the area

The studied areas are located in the experimental area of the State University of Southwest of Bahia, Campus of Vitória da Conquista, at the geographic coordinates of 14°53' latitude south and 40°48' longitude west. The climate of the region is subtropical highland (Cwb) according to the Köppen classification, with an annual average temperature of 21 °C and precipitation between 700 mm and 1100 mm. The region has an altitude of around 880 m. The soils of the studied areas have a clayey texture and belong to the single class of Dystrophic Yellow Latosol (Santos et al., 2006). Table 1 shows the chemical characterization of these soils at a depth of 0-10 cm.

Table 1
Chemical characterization of the soil (0-10 cm depth).

Cover	pH	OM	P	K	Ca	Mg	H+Al	Al	V	M
		g dm–3	mg dm–3	————— cmolc dm3 —————					—— % ——
Pinus in consortium*	5.1	20	1.0	0.23	1.0	0.9	3.8	0.3	36	12
Pinus caribaea	4.7	18	1.0	0.14	1.0	1.0	4.8	0.8	27	31
Native forest	4.9	36	3.0	0.17	3.2	1.4	5.8	0.3	45	6

• ^*
  Pinus in consortium – Pinus caribaea, Pinus oocarpa, Pinus taeda; RS = Reproductive structures; Analyses performed according to EMBRAPA (1997): pH (water); extractable P and K by Mehlich-1; exchangeable Ca, Mg and Al by KCl 1 mol L^–1 and organic matter (OM) by oxidation with Na₂Cr₂O₇ 4 N. Soil samples were collected at a depth of 0-10 cm, with four replicates, one per each 15 × 15 m plot. Each replicate consisted of a sample composed of 20 simple samples. V-base saturation and m-saturation by aluminum.

The study was conducted in two Pinus stands, one in consortium (composed of three species: Pinus caribaea, Pinus oocarpa, and Pinus taeda), and another stand purely composed of Pinus caribaea var. hondurensis. Each stand has an area of approximately 0.5 ha and is about 11 years old. A fragment of Semi-deciduous Seasonal Forest, regionally known as mata-de-cipó, was used as reference.

The Pinus stands were implanted in an area of lower capoeira vegetation after the area had been cleared using a crawler tractor and the soil harrowed. Planting was carried out in 2002 with seminal seedlings and pit fertilization (200 grams of Simple Superphosphate), according to a 3 m × 3 m spacing. Stand maintenance was carried out at two, six and twelve months after planting, with weed-competition control by weeding the lines and between the rows. According to measurements made in June 2013 in the Pinus stand consortium, the diameter at breast height (DBH) of the trees varied from 13.2 to 25.0 cm for Pinus oocarpa, between 13.8 and 25.5 cm for Pinus caribaea, and between 5.0 and 13.7 cm for Pinus taeda. In pure stand, the Pinus caribaea trees presented a DBH between 11.9 and 23.9 cm.

The reference fragment has an area of 42 hectares and is in the middle stage of ecological succession according to criteria established by Brasil (1994). The vegetation consists of partially deciduous woody plants surrounded by lianas, with height varying between 10 and 20 m and an ecotype predominance of the Leguminosae family, with the genus Parapiptadenia being highlighted (IBGE, 2012).

2.2. Litterfall production

Wooden collector trays of 0.5 m × 0.5 m (0.25 m²) suspended 15 cm away from the soil surface and with 1 mm nylon mesh bottom were used. Twelve (12) collectors were randomly distributed in each of the Pinus stands in four 15 m × 15 m plots (three collectors per plot), which were also randomly demarcated. A 1,000 m² (100 × 100 m) plot was established in the Semi-deciduous Seasonal Forest fragment where four collectors were randomly installed, as described by Gama-Rodrigues & Barros (2002).

The litter intercepted by the collectors was collected monthly during twelve months (from June 2013 to May 2014). The litter was sorted after each collection into: leaves, reproductive structures, bark and branches. After sorting the samples, the fractions were oven dried (at 60 °C for 72 hours) and then weighed.

Based on the dry mass results of the fractions, the monthly and total production of forest ecosystems were calculated according to the following formulas:

$Pro{d}_{month/ha}=\frac{\left(Pro{d}_{month}*10.000\right)}{A_C}$ $Pro{d}_{total/ha}=\sum Pro{d}_{month/ha}$

In which: Prod_month = Litterfall contribution obtained in each month (kg month^–1); Prod_month/ha = Monthly contribution of litterfall per hectare (kg ha^–1 month^–1); Ac = Collector Area (m²); Prod_total/ha = Total amount of litterfall (kg ha^–1).

2.3. Accumulated litterfall

The amount of litterfall accumulated on the soil surface was estimated using a 0.5 × 0.5 m (0.25 m²) square wooden frame, which was randomly launched with four replicates in each area. Three collections were performed: one in the first month of the experiment, the second six months after the beginning, and the last at the end of the twelve months. All deciduous plant material deposited on the soil and circumscribed to the frame at different degrees of decomposition was considered as accumulated litter. The same drying and weighing procedure used for the estimation of litter deposited on the collectors was adopted.

2.4. Decomposition rate

The equation proposed by Olson (1963) was used to estimate the litter decomposition rate: K = L/X, in which K = coefficient of decomposition in the condition of dynamic equilibrium, L = annual litterfall production (kg ha^–1), and X = annual average accumulated litter (kg ha^–1). Based on the K value, the mean renewal time (T_R) of accumulated litter on the soil was estimated by the relationship 1/K, and the half-life (T_0.5); moreover, the time required for 50% litter decomposition to occur was found using the equation T_0.5 = –Ln 0.5/K, as proposed by Shanks & Olson (1961).

2.5. Climate variables

The climatic variables of precipitation, temperature, wind speed and relative humidity were considered to assess the influence of climate on litterfall production. The data corresponding to the study period (from June 2013 to May 2014) were made available by the Meteorological Station of the State University of Southwest of Bahia (ESMET), as shown in Table 2.

Table 2
Climatic data of the study period in Vitória da Conquista, Bahia state, Brazil. Data provided by the Meteorological Station of the State University of Southwest Bahia (ESMET).

Months	Climate Variables
	Precipitation	Average temperature	Wind speed	Relative humidity
	mm	°C	m s–1	%
June 2013	35.30	19.78	1.71	87.30
July 2013	17.10	19.36	1.98	84.35
August 2013	19.30	19.46	2.59	80.69
September 2013	20.80	21.01	2.61	78.13
October 2013	29.30	21.68	2.82	75.09
November 2013	33.20	23.10	2.51	69.13
December 2013	231.70	23.06	1.55	77.72
January 2014	69.60	22.17	2.45	76.57
February 2014	35.30	20.81	2.50	77.25
March 2014	37.00	23.00	2.03	75.04
April 2014	42.00	22.94	2.20	71.78
May 2014	42.00	21.67	1.63	72.51

2.6. Statistical analyses

The data were submitted to normality analysis (Lilliefors test) and homogeneity of variance of error (Cochran and Bartlett test). Analysis of variance according to a completely randomized design with four replications was performed for parametric data, while the Tukey test at 5% significance was adopted to compare means between forest ecosystems and litter fractions.

In order to evaluate the influence of climatic factors on litterfall production over the study period, Pearson correlations at 5% significance were established between litter deposition and climatic variables (precipitation, temperature, wind speed and relative humidity). All statistical analyses were carried out using the statistical program SAEG^® v.9.1.

3. RESULTS AND DISCUSSION

3.1. Litterfall production

The average monthly litterfall production of Pinus stands was 213.7 kg ha^–1 month^–1. This value is similar to that observed by Vieira & Shumacher (2010a) in a pure stand of Pinus taeda at 11 years of age in Cambará (RS) (212.1 kg ha^–1 month^–1); however, it is lower than that found by Melo & Resck (2002) in studying Pinus caribaea stands at 16 years of age in the Cerrado region, who found a deposition rate of 868.25 kg ha^–1 month^–1 for the hondurensis variety, and 596.8 kg ha^–1 month^–1 for the bahamensis variety. Poggiani (1987) also verified a higher result in plantations of the Pinus genus at 11 years of age in Agudos (SP), where contributions of 700.0kg ha^–1 month^–1 for Pinus caribaea var. hondurensis, and of 591.7 kg ha^–1 month^–1 for Pinus oocarpa were found.

The average monthly litter contribution of the native forest was 527.4 kg ha^–1 month^–1. This result is similar to that observed by Santos Neto et al. (2015) in the Semi-deciduous Seasonal Forest located in the same municipality where the present study was carried out (544.6 kg ha^–1 month^–1). On the other hand, Pinto et al. (2008) found a higher value in a Semi-deciduous Seasonal Forest in Viçosa (MG) (735.0 kg ha^–1 month^–1).

In comparing the average monthly production of total litterfall between the studied coverages, it was observed that Pinus stands did not differ from each other, and presented lower values than those observed in the native forest (Table 3). This result indicates that the organization of the forest system (homogeneous or heterogeneous) did not provide differences in the litterfall deposition dynamics of Pinus stands. The same tendency was observed for the monthly depositions of leaves, bark and branches. Hinkel & Panitz (1999) also found a higher monthly contribution in the native forest area (Mata Atlântica) (530.88 kg ha^–1 month^–1) when compared to a Pinus elliottii stand (354.17 kg ha^–1 month^–1) in Florianópolis (SC).

Table 3
Average monthly litter production in a twelve-month period in three forest ecosystems in southwestern Bahia.

Coverage	Litter Fractions (kg ha–1 month–1)
	Leaves	RS	Bark		Branches		Total
Pinus in consortium*	225.7 (12.9) Ab	5.0 (1.5) Bb	2.0 (0.3) Cb	1.5 (1.4) Cb		234.2 (13.2) b
Pinus caribaea	193.0 (13.2) Ab	0.4 (0.4) Bb	0.7 (0.3) Bb	1.3 (0.9) Bb		195.4 (13.4) b
Native forest	381.2 (16.4) Aa	14.7 (4.6) Ca	30.2 (9.6) Ca	101.3 (11.8) Ba		527.4 (24.7) a

• ^*
  Pinus in consortium – Pinus caribaea, Pinus oocarpa, Pinus taeda; RS = Reproductive structures. Values in parentheses correspond to the mean standard error. Averages followed by the same uppercase letters in the line which compare the fractions, and lowercase letters in the column which compare forest ecosystems, do not differ from one another by the Tukey test at 5% significance.

Total annual litterfall for Pinus in the consortium, Pinus caribaea and native forest were 2.780, 2.406 and 6.497 kg ha^–1, respectively (Table 4). Novais & Poggiani (1983) observed higher results in Pinus plantations in a homogeneous stand of Pinus caribaea (4.458 kg ha^–1) and in a consortium of Pinus caribaea with Liquidambar (5.571 kg ha^–1) in the municipality of Agudos (SP). However, in studying a Semi-deciduous Seasonal Forest in a late successional stage in the state of Minas Gerais, Pinto et al. (2008) found annual production of 8.819 kg ha^–1, a value higher than that observed in the reference forest of the present study.

Table 4
Monthly production and contribution of different litter forming fractions in a twelve-month period in three forest ecosystems in Southwest Bahia.

Month/ Year	Litter Fractions
	Leaves		RS		Bark		Branches		Total
	kg·ha–1	%	kg·ha–1	%	kg·ha–1	%	kg·ha–1	%	kg·ha–1
	Pinus in consortium *
June	384.6	97.99	6.44	1.64	0.87	0.22	0.58	0.15	392.44
July	168.1	91.13	11.85	6.43	4.51	2.44	0.00	0.00	184.44
August	188.5	76.94	55.64	22.71	0.84	0.34	0.00	0.00	244.95
September	234.0	95.52	9.78	3.99	1.20	0.49	0.00	0.00	245.02
October	267.4	97.25	6.51	2.37	1.05	0.38	0.00	0.00	274.98
November	307.9	97.88	4.69	1.49	1.96	0.62	0.00	0.00	314.51
December	215.0	97.32	2.04	0.92	3.89	1.76	0.00	0.00	220.95
January	164.6	96.10	0.56	0.33	6.12	3.57	20.40	11.91	191.88
February	104.0	98.21	0.33	0.31	0.84	0.79	0.73	0.00	105.93
March	188.8	99.03	0.04	0.02	1.82	0.95	0.00	0.00	190.62
April	255.5	99.59	0.95	0.37	0.11	0.04	0.00	0.00	256.51
May	156.9	99.17	0.22	0.14	1.09	0.69	0.00	0.00	158.22
Total	2635.2	94.66	99.03	3.39	24.30	0.99	21.71	0.96	2780.23
	Pinus caribaea
June	222.6	99.30	1.03	0.46	0.53	0.24	0.00	0.00	224.13
July	215.4	98.73	1.70	0.78	1.07	0.49	0.00	0.00	218.13
August	135.7	95.41	0.67	0.47	0.20	0.14	5.67	3.98	142.23
September	192.3	99.59	0.63	0.33	0.17	0.09	0.00	0.00	193.07
October	288.9	99.85	0.30	0.10	0.13	0.05	0.00	0.00	289.30
November	243.8	99.61	0.17	0.07	0.80	0.33	0.00	0.00	244.80
December	213.6	99.49	0.50	0.23	0.60	0.28	0.00	0.00	214.70
January	108.5	99.97	0.03	0.03	0.00	0.00	0.00	0.00	108.53
February	79.6	98.16	0.07	0.08	0.87	1.07	0.57	0.70	81.13
March	145.4	99.75	0.13	0.09	0.23	0.16	0.00	0.00	145.77
April	368.4	100.00	0.00	0.00	0.00	0.00	0.00	0.00	368.43
May	162.5	92.43	0.00	0.00	3.77	2.14	9.53	5.42	175.80
Total	2376.7	98.52	5.23	0.22	8.37	0.41	15.77	0.84	2406.03
	Native forest
June	423.2	77.62	47.50	8.71	0.00	0.00	74.50	13.66	545.20
July	441.9	76.40	70.80	12.24	0.00	0.00	65.70	11.36	578.40
August	699.4	78.06	24.40	2.72	17.90	2.00	154.30	17.22	896.00
September	556.5	75.74	0.00	0.00	84.80	11.54	93.47	12.72	734.80
October	835.2	68.87	0.00	0.00	130.80	10.79	246.67	20.34	1212.67
November	194.0	75.86	0.13	0.05	2.40	0.94	59.20	23.15	255.73
December	507.9	87.99	0.00	0.00	2.00	0.35	67.33	11.67	577.20
January	114.5	58.39	2.60	1.33	5.40	2.75	73.60	37.53	196.10
February	132.1	27.21	0.40	0.08	64.80	13.35	288.20	59.36	485.50
March	152.8	75.79	8.50	4.22	0.90	0.45	39.40	19.54	201.60
April	217.0	87.01	7.10	2.85	0.10	0.04	25.20	10.10	249.40
May	442.5	78.47	0.00	0.00	78.30	13.89	43.10	7.64	563.90
Total	4717.0	72.28	161.43	2.68	387.40	4.67	1230.67	20.36	6496.50

• ^*
  Pinus in consortium – Pinus caribaea, Pinus oocarpa, Pinus taeda; RS = Reproductive structures.

The leaves fraction represented the greatest proportion of the litterfall in both the Pinus stands and the native forest for all months of the year (Table 4), thus contributing an average of 96.6% and 72.2% to the litterfall, respectively (Table 3). The mean value of Pinus plantations agrees with those found by Piovesan et al. (2012) (95.6%) in Pinus taeda stands in the state of Paraná. Several other studies in natural and planted forests have also found that the leaves fraction constitutes the main component of the material deposited on the soil (Schumacher et al., 2003; Pinto et al., 2008; Diniz et al., 2011; Cunha et al., 2013; Antoneli & Schenemann, 2014; Santos Neto et al., 2015).

The average contribution of the other fractions in the Pinus caribaea and native forest stands followed the order: leaves > branches > bark > reproductive structures. Pezzatto & Wisniewski (2006) found the same distribution pattern in different succession species of the Semi-deciduous Seasonal Forest in the West of Paraná, and Piovesan et al. (2012) in Pinus taeda stands. The distribution sequence in the consortium Pinus stand was: leaves > reproductive structures > bark > branches.

Based on Table 4, which shows the variation of the total litter production and the fractions throughout the year, it is observed that there was no production of branches in the Pinus stands for most months. According to Collins (1977), the use of collectors can lead to faults in the sampling of the branches fraction, considering that they might not reach the collectors. However, in view of the fact that the branches in Pinus stands are better adhered to the tree trunks when compared to native forest, this may explain the null production of these tree branches in several months of the year.

3.2. Temporal variation of litterfall production

The maximum contribution of total litter occurred in June for the Pinus consortium stand, and in April for Pinus caribaea (Table 4). The maximum contribution in the native forest occurred in October, with a decrease after this month (Table 4). Such distribution characterizes a typical seasonal pattern of semi-deciduous seasonal forests, in which the peak leaf deposition occurs at the end of the dry season (August-October) as the vegetation’s response to the climatic seasonality (Werneck et al., 2001; Arato et al., 2003; Santos Neto et al., 2015).

When the data regarding the monthly litter production (total and fractions) was correlated with the climatic variables, significant associations were only found between the branches fraction and the wind speed in the native forest; between the reproductive structures (RS) fraction with relative humidity (positive correlation) in the native forest and in the Pinus in consortium; and with temperature (negative correlation) in all forest ecosystems studied (Table 5). These results indicate that the largest deposition of the reproductive structures occurred during the period of lower temperatures and higher humidity. This observation is consistent with a study carried out by Piovesan et al. (2012) in Pinus taeda in the state of Paraná; by König et al. (2002) in Deciduous Seasonal Forest in Rio Grande do Sul, who found negative coefficients between temperature and the production of reproductive structures; and by Vieira & Shumacher (2010a) for Pinus taeda in Cambará do Sul (RS), who found a positive correlation between the deposition of reproductive structures and relative humidity.

Table 5
Correlation coefficient between the monthly variation of litter production and meteorological variables.

Climate Variables	Litter Fractions
	Leaves	RS	Bark	Branches	Total
	Pinus in consortium
Precipitation	0.10	-0.29	0.45	0.09	0.09
Average temperature	0.15	-0.66*	0.11	0.13	0.11
Wind speed	-0.20	0.12	-0.15	0.17	-0.19
Relative humidity	0.08	0.61*	0.15	-0.01	0.12
	Pinus caribaea
Precipitation	-0.04	-0.25	0.34	0.09	-0.09
Average temperature	0.47	-0.56*	0.02	0.14	0.36
Wind speed	0.03	0.37	-0.13	0.19	0.10
Relative humidity	-0.49	0.29	0.12	-0.03	-0.43
	Native forest
Precipitation	0.01	-0.27	-0.25	-0.18	-0.10
Average temperature	-0.42	-0.72*	-0.11	-0.33	-0.48
Wind speed	0.15	-0.30	0.44	0.58*	0.32
Relative humidity	0.30	0.78*	-0.19	0.11	0.29

• ^*
  significant correlations at 5% significance.

3.3. Rate of litter decomposition

The average amounts of accumulated litter observed in Pinus stands were lower than the value found in native forest (Table 6). Higher accumulations were verified by Carvalho et al. (2008) in Pinus elliottii stands (18.000 kg ha^–1), and by Lima et al. (2015) in Pinus oocarpa stands (15.267 kg ha^–1) over 30 years of age.

Table 6
Decomposition constant (K), litter renewal time and litter half-life (T_1/2) of three forest ecosystems in southwestern Bahia.

Cover	Accumulation	K	TR	T0.5
	kg ha–1		---- years ----
Pinus in consortium*	3590.95 (215.3) b	0.77 (0.09) a	1.29 (0.13)	0.90 (0.09)
Pinus caribaea	2822.40 (207.0) b	0.85 (0.37) a	1.17 (0.12)	0.81 (0.08)
Native forest	6385.20 (399.0) a	1.02 (0.12)a	0.98 (0.15)	0.68 (0.11)

• ^*
  Pinus in consortium – Pinus caribaea, Pinus oocarpa, Pinus taeda; K = decomposition constant; T_R = litter renewal time; T_0.5 = half-life time. Values in parentheses correspond to the mean standard error. Means followed by the same letter in the column do not differ from one another by the Tukey test at 5% significance.

The coefficient of litter decomposition (K) did not significantly vary among the studied stands (Table 6), which shows that they are not distinguishable as to the decomposition speed of their vegetation residues. However, low decomposition rates are usually observed in Pinus stands (Melo & Resck, 2002; Ribeiro et al., 2007; Carvalho et al., 2008), which are generally attributed to the chemical composition of the plant material and presents high levels of phenolic compounds and lignin and/or a higher C/N ratio (Moreira & Siqueira, 2002).

Morellato (1992), Schlittler et al. (1993) and Pimenta et al. (2011) recorded K coefficients ranging from 1.02 to 2.45 in fragments of Semi-deciduous Seasonal Forest. In addition to the quality of plant residues, variations in the litter decomposition speed among forest stands may be related to the type of vegetation cover, to the fauna activity of the soil (Anderson et al., 1983) and to environmental conditions (Silva et al., 2014).

The average time for litter renewal (T_R) in Pinus stands was 1.2 years (Pinus caribaea) and 1.3 years (Pinus in consortium), while it was 0.98 years in the native forest (Table 6). According to Vogt et al. (1986) and Melo & Resck, (2002), T_R values in tropical deciduous and semi-deciduous hardwood forests are usually observed as being less than one year, while this time is greater than one year in the case of conifers (Pinus).

The time required for 50% litter decomposition (T_1/2) to occur was estimated at 0.90 years (329 days) for Pinus in consortium, 0.81 years (296 days) for Pinus caribaea, and 0.68 years (248 days) for the native forest. Cunha et al. (2013) and Lopes et al. (2009) found T_1/2 of 0.73 and 0.98 years for Semi-deciduous Seasonal Forest in secondary succession in the state of Minas Gerais and for the Caatinga in Ceará, respectively.

4. CONCLUSION

1. ¹

   Litterfall production is similar for both Pinus stands, which allows for inferring that the organization of the forest system (homogeneous or heterogeneous) did not interfere in litterfall contribution dynamics. On the other hand, these values are inferior to those found in the reference forest and in stands of the same genus in other regions of Brazil, thus showing lower potential for litter accumulation.

2. ²

   The litter produced was mostly composed of leaves; about 97% in the Pinus stands, and 72% in the native forest.

3. ³

   The temporal variation of litter supply is not very sensitive to variations in climate, with a greater influence of the temperature and relative air humidity in the deposition of reproductive structures.

4. ⁴

   The litter decomposition speed is similar for the Pinus stands and the native forest. However, the average litter renewal time is over one year in Pinus stands (mean of 1.25 years), and less than one year in the native forest (0.98 years).

• FINANCIAL SUPPORT FAPESB, bolsa de iniciação científica, n° de outorga: 1925 /2013.

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Publication Dates

• Publication in this collection
  2 May 2019
• Date of issue
  2019

History

• Received
  16 Apr 2016
• Accepted
  26 Oct 2017