Journal of Sustainable Forestry

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Effect of Thinning Intensity on Litterfall

Biomass and Nutrient Deposition in a Naturally
Regenerated Pinus Pseudostrobus Lind. Forest in
Oaxaca, Mexico

Yazmin Pérez-Alavez, Gerardo Rodríguez-Ortiz, Wenceslao Santiago-García,

Gisela Virginia Campos-Angeles, José Raymundo Enríquez-del Valle &
Meredith P Martin

To cite this article: Yazmin Pérez-Alavez, Gerardo Rodríguez-Ortiz, Wenceslao Santiago-García,

Gisela Virginia Campos-Angeles, José Raymundo Enríquez-del Valle & Meredith P Martin (2021):
Effect of Thinning Intensity on Litterfall Biomass and Nutrient Deposition in a Naturally Regenerated
Pinus�Pseudostrobus Lind. Forest in Oaxaca, Mexico, Journal of Sustainable Forestry, DOI:
10.1080/10549811.2021.1946410

To link to this article: https://doi.org/10.1080/10549811.2021.1946410

Published online: 26 Jul 2021.

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JOURNAL OF SUSTAINABLE FORESTRY
https://doi.org/10.1080/10549811.2021.1946410

Effect of Thinning Intensity on Litterfall Biomass and Nutrient

Deposition in a Naturally Regenerated Pinus Pseudostrobus
Lind. Forest in Oaxaca, Mexico
Yazmin Pérez-Alaveza, Gerardo Rodríguez-Ortiza, Wenceslao Santiago-García b
,
Gisela Virginia Campos-Angelesa, José Raymundo Enríquez-del Valle a,
and Meredith P Martin c
a
División de Estudios de Posgradoe Investigación (DEPI), Tecnológico Nacional De México-Instituto Tecnológico
Del Valle De Oaxaca, Xoxocotlán, Oaxaca, Mexico; bInstituto de Estudios Ambientales-División de Estudios de
Postgrado, Universidad De La Sierra Juárez, Camino a La Universidad S/N, Ixtlán De Juárez, Oaxaca, Mexico;
c
Department of Forestry and Environmental Resources, North Carolina State University, Raleigh, North
Carolina, USA

ABSTRACT KEYWORDS
Litterfall is an important component of forest biomass and nutrient Leaf litter; carbon; nitrogen;
cycling, and can have key impacts on soil fertility through its decom­ biomass; thinning intensity;
position. However, the effect of forest management on litterfall silviculture; pine
remains unclear. We evaluate the impact of thinning intensity on the
biomass and nutrient content (C and N) of litterfall in Pinus pseudos­
trobus forest in Oaxaca, México, across two thinning intensities in areas
with either high or low residual basal area and across seasons. There
was significantly higher litter carbon content in the least intensive
thinning treatment, but no significant differences in biomass or nitro­
gen content between treatments. However, there was a significant
correlation between residual basal area and litter biomass at the p < .1
level. We found a clear seasonal pattern in litterfall fluxes, with 1.5
times more litter deposition in autumn’s dry season compared to
spring. We find that the thinned stand of P. pseudostrobus generated
an annual mean litter biomass of 1059.27 ± 346.04 kg ha−1 with mean
carbon content of 125.31 ± 46.43 kg ha−1 and mean nitrogen content
of 4.76 ± 1.43 kg ha−1. These values present an important contribution
for modeling of biomass and nutrient cycling in this ecologically and
economically important forest type.

Introduction
Carbon sequestration in forests has generated widespread international interest due to its
potential to mitigate climate change (Bonan, 2008; Breedlow et al., 2004). While much
attention is paid to understanding and estimating carbon capture within forest woody
biomass, tree litterfall also contains high levels of carbon and mineral nutrients. On both
stand and global levels, litterfall represents between 4–6% of all forest carbon content (Pan
et al., 2011; Rodríguez-Ortiz et al., 2012). In addition, foliage is one of the most important
components of stand productivity, as leaves are the site of photosynthesis and therefore of
a forest’s primary productivity. Furthermore, the nutrients contained in the foliage can be

CONTACT Meredith P Martin mpmarti7@ncsu.edu Department of Forestry and Environmental Resources, North
Carolina State University, Raleigh, North Carolina 27695, USA
© 2021 Taylor & Francis
2 Y. PÉREZ-ALAVEZ ET AL.

considerable, particularly for the macro and micronutrient minerals essential to plant
growth (González-Rodríguez et al., 2019; Meentemeyer et al., 1982). Because of this, the
decomposition of litter on the forest floor is an important mechanism for conserving soil
fertility, increases the productive capacity of a site, and is integral to forest ecosystem
functioning (Attiwill & Adams, 1993; Roig et al., 2005; Vitousek, 1984). In fact, in some
forest systems with low soil fertility, the availability of nutrients appears to be determined by
the amount of organic matter left for decomposition from litterfall and from slash left from
harvest operations (Merino et al., 2003; Zianis & Mencuccini, 2003).
It is essential to understand the rates of leaf-litter fall and decomposition to organic
material for sustainable management, and especially to understand the relationship between
these factors and stand structure. The rate of litterfall and decomposition directly impacts site
productivity, which in turn affects processes as diverse as photosynthesis, soil erosion, and the
production of both timber and non-timber forest products (Kunhamu et al., 2009; Roig et al.,
2005). Different silvicultural treatments such as harvests, thinnings, and pruning generate
large quantities of plant debris which is often left on the ground. These treatments also impact
the residual stand density and basal area, which in turn impact the amount of leaf litter
generated (Berg & Meentemeyer, 2001; Blanco et al., 2006; Bueis et al., 2018; Çömez et al.,
2019; Huebschmann et al., 1999; Jiménez & Navarro, 2016; Kim, 2016; Navarro et al., 2013).
The majority of studies examining the impact of silvicultural management on litterfall
dynamics focus on temperate forests in Europe, and little research exists in other parts of the
world. There is, therefore, a need for research on litterfall dynamics in Mexico, where
timber production is a major source of income (SEMARNAT, 2020). Nationally, pine
represents 75% of all wood production in Mexico, with an economic value of over
7 billion pesos in 2016 (SEMARNAT, 2016). In Oaxaca state, pine represents approximately
80% of all timber production, and generated almost 400 million pesos in 2016
(SEMARNAT, 2016). Within Mexico, Pinus pseudostrobus Lindl. is a widespread timber
species with high potential for use in commercial forestry plantations, as the tree has high
quality wood, a fast growth rate, and is a good resin producer (Cambrón-Sandoval et al.,
2014, 2013; López-Upton, 2002; Sáenz-Romero et al., 2012). In the community of San Pedro
el Alto in Oaxaca state, P. pseudostrobus represents 15% of the 100,000 m3 of wood
harvested each year (Servicios Técnicos Forestales (STF), 2014).
However, little research examines the nutrient cycling or litterfall dynamics in either
natural forest or plantation systems for this or other pine species in southwestern Mexico.
This study examines the impact of different intensities of thinning treatments on litterfall
biomass and nutrient content in natural P. pseudostrobus forest in this community forest in
Oaxaca, México. With this research, we provide a key contribution to the broader under­
standing of seasonal nutrient dynamics and the impacts of management on nutrient cycling
in these ecologically and economically important systems.

Materials and methods

Study area and stand measurements
This study examined two thinning treatments applied to a pine stand in San Pedro el Alto in
the Sierra Sur region of Oaxaca, Mexico. Experimental thinning treatments were established
in 2018 in a young stand in stem exclusion phase located at a mean altitude of 2800 m.a.s.l.
 JOURNAL OF SUSTAINABLE FORESTRY 3

at 16° 80´ 83,33 N and −097° 05´ 33,3” W. The mean annual temperature is 18°C with mean
total annual precipitation of 1500 mm. The climate is seasonally dry, temperate, with a rainy
season in the summer (generally from May or June to October or November) when most of
this precipitation falls (CONAGUA, 2020). The forest type is dominated by Pinus pseudos­
trobus, with associated trees of P. oaxacana Mirov, P. patula Schl. et Cham., several oak
species (Quercus sp.), Cedrus sp., and understory trees and shrubs such as madrone
(Arbutus sp.) and blackberry (Rubus sp.), although this particular regenerating stand was
composed of pure P. pseudostrobus at the time of this study. The P. pseudostrobus here
naturally regenerated following a fire, and consisted of a single cohort of 10–12 years in age.
The stand (about 20 ha in size) was thinned in 2016 at two levels of intensity: a high-
intensity thinning removed 75% of stems, and a low-intensity thinning removed 50% of
stems. Both thinning treatments took the form of pre-commercial low thinnings conducted
by the local community, where trees with smaller diameters or poor form were removed. In
2018, we established experimental circular sites of 400 m2 using a random design of site
location based on two variables: thinning intensity and residual basal area, but ensuring that
the site location fell at least 100 m from the edge of each treatment type. Each of these two
variables had two treatment levels of either high- or low-thinning intensity (50 or 75% of
pre-thinning tree density removed) or high or low residual basal area (H for high basal area
> 13 m2 ha−1; L for low basal area ≤ 13 m2 ha−1). These four treatments are termed H50,
H75, L50, L75 from here on, and contained three replicates each for a total of 12 plots. In
each plot, we measured all trees present for diameter (DBH) with a Haglof caliper for post-
thinning basal area and density calculations. Pre-thinning densities at each site were

Figure 1. Photos of the field sites showing (a) an example mesh litter trap, (b) a site in more intensively
thinned area with lower residual basal area, and (c) a site in a less intensively thinned area with higher
residual basal area.
4 Y. PÉREZ-ALAVEZ ET AL.

Table 1. Summary of stand structure characteristics at each sampling site with pre- and
post-thinning tree density. Mean DBH is shown in cm, density in trees ha−1 and basal area
in m2 ha−1. Thinning level represents the thinning intensity (50% or 75% of stems
removed) and the residual basal area as high or low (H for >13 m2 ha−1 and L for
≤13 m2 ha−1).
Density Density Mean DBH Basal Area Thinning
Site
(pre-thinning) (post-thinning) (post-thinning) (post-thinning) level

1 1800 900 15.42 18.35 50H

2 2000 500 20.28 24.4 75H
3 2900 725 15.51 22.38 75H
4 1350 338 18.13 15.4 75H
5 1200 600 17.77 12.25 50L
6 650 325 18.21 10.71 50L
7 1375 344 15.00 9.91 75L
8 1300 325 17.32 12.84 75L
9 2145 536 15.07 8.96 75L
10 1475 738 20.57 24.53 50H
11 1000 500 21.43 14.22 50H
12 1400 700 16.62 9.57 50L

estimated based on the number of stumps left (see Figure 1 for example, photographs of the
sites). For a summary of stand structure pre- and post-thinning at each site, see Table 1.

Litterfall sampling and analysis

In each circular site we established five leaf-litter traps 60 cm above the soil (one in each
quadrant and one in the center) in October 2018 follow a systematic design. The sites were
located inside a 100 m buffer from the edge of each thinning treatment to ensure that all
litterfall caught in our traps was from the appropriate treatment. Within each site, we
systematically located litter traps with one in the center of the circular site and the other four
located at the four cardinal directions from the center. We used circular traps, measuring
0.725 m in diameter and made of wire and mesh (greenhouse shade cloth with 64
cells cm−1). Traps were checked every month for a year, and all litter that fell in the five
traps was collected as a composite by site for analysis (including foliage, twigs, male flowers,
and bark). The litter was composed of components solely from P. pseudostrobus, as other
species were not present within our sample sites. The litterfall were dried at 75°C for 120 h
in a laboratory convection oven (Memmert Gmbh +Co. KG) to obtain dry weight, mea­
sured with a Sartorius analytic balance scale. Total seasonal and annual litterfall biomass
were also calculated and recorded.
To calculate carbon and nitrogen content, composite samples of approximately 700 g
were taken from accumulated biomass from each site in each month. Litterfall from the five
traps were thoroughly mixed together with all components included (i.e. leaves, bark, twigs,
flower parts), and a sample was taken from this mixture such that the proportions of litter
components were maintained within the sample. These samples were ground (with all litter
components together) and then oven-dried until constant weight at a temperature of 70°C.
 JOURNAL OF SUSTAINABLE FORESTRY 5

Carbon concentration (C) was determined using the Walkley and Black method, and
nitrogen concentration (N) was determined using the Micro-Kjendahl method following
standard protocols of NOM-021-SEMARNAT-2000 (NOM-021-SEMARNAT-2000, 2002;
Ma & Zuazaga, 1942; Walkley & Black, 1934).

Data analysis
All statistical analyses were performed using the program SAS (Statistical Analysis System
(SAS), 2015). To analyze monthly litter biomass and nutrient fluxes (C and N per 700 g of
litter) by thinning intensity and seasonality, we used a two-way ANOVA using the PROC
GLM model and tested pair-wise significant differences with a Tukey test on monthly
deposition values (with p < .05 significance level). We also estimated the mean annual litter
biomass and N and C content on a per hectare basis in this forest type. To meet assumptions
of normality and homogeneity of variance based on the Shapiro Wilk and Bartlett tests, we
pffiffi
transformed the nutrient variables using a log 10 xtransformation for all analyses. To
assess changes in litterfall across seasons, we defined spring as March 20th – June 21st,
summer as June 22nd – September 21st, autumn as September 22nd – December 20th, and
winter as December 21st – March 19th. These seasons are slightly different from standard
temperate definitions as they are based on a combination of temperature changes and
changes between the rainy and dry seasons in Oaxaca.
We used a cluster analysis to classify experimental sites based on their transformed
annual nutrient concentrations (C and N) and annual accumulated biomass using mean
square Euclidean distance and hierarchical clustering. We then examined the stand char­
acteristics of the resulting clusters to assess whether there were any clear patterns. Finally,
we performed linear regressions between biomass and residual basal area and residual tree
density, and between annual nutrient content (log10 transformed) and residual basal area
and residual tree density to more directly examine the relationship between current stand
structure and litterfall deposition.

Results
Effects of thinning treatment and season on litterfall
The litterfall in this stand of Pinus pseudostrobus generated 1059.27 kg ha−1 of mean annual
litter biomass and 125.32 kg ha−1 of carbon and 4.76 kg ha−1 of nitrogen. Overall, carbon and
nitrogen represented 11.75% and 0.40% of litterfall biomass. We found distinct seasonal
patterns in litterfall with two peaks in biomass, carbon and nitrogen (Figure 2). A sustained
peak in litter biomass occurred first in the fall months from October-December, followed by the
lowest litter deposition in summer months, and then another small peak in litterfall in March.
In the analysis of variance of season and thinning intensity on litter nutrient and biomass
content, there were significant differences in carbon content by thinning intensity but not in
biomass or nitrogen (Table 2; two-way ANOVA, p < .05). Biomass, N and C all showed
significant differences between seasons, but the interaction between season and thinning
intensity was not significant (Table 2; two-way ANOVA, p < .01).
Mean biomass of P. pseudostrobus litterfall across all treatments and seasons was
256.60 kg ha−1. The highest mean litter biomass was found in sites under the H50 treatment
 6
Y. PÉREZ-ALAVEZ ET AL.

Figure 2. Litterfall biomass, carbon, and nitrogen content (kg) by month for one year (October 2018 – September 2019) across the four thinning treatments as
mean and standard deviation. The four treatments represent either 50% or 75% of basal area removed for thinning intensity, and H for high residual BA (˃13 m2
ha−1) or L for low (≤13 m2 ha−1).
 JOURNAL OF SUSTAINABLE FORESTRY 7

Table 2. Summary of two-way ANOVA results of the effect of thinningpintensity

ffiffi and season on biomass,
carbon, and nitrogen. Both carbon and nitrogen variables are log 10 x transformed. Asterisks indicate
statistical significance: “**” for p < .01, “*” for p < .05.
Predictor Variables Degrees of freedom Response variables
Biomass Carbon Nitrógen
(kg ha−1) (kg ha−1) (kg ha−1)
Thinning intensity 3 18,375.231 0.021* 0.009
Season 3 31,362.595** 0.025** 0.025**
Thinning×Season 9 2507.852 0.002 0.002
Error 32 10,247.751 0.006 0.006
Total 47

Table 3. Mean annual biomass and total amounts of nutrients deposited by litterfall across thinning
treatments, and mean monthly biomass and nutrient content across seasons. Thinning treatments are
based on a combination of residual basal area (H for >13 m2 and L for ≤ 13 m2) and thinning intensity
(50% or 75% of stems removed in thinning). Letters represent significant pair-wise differences based the
Tukey test (p < .05), and error represents standard deviation.
Thinning Treatment Biomass (kg ha−1) Carbon (C) (kg ha−1) Nitrogen (N) (kg ha−1)
H50 1280.11 ± 603.06 166.45 ± 78.45a 5.34 ± 2.51
H75 1004.37 ± 117.85 115.73 ± 13.58ab 4.58 ± 0.54
L50 1009.05 ± 393.50 106.20 ± 41.41b 4.05 ± 1.58
L75 943.53 ± 156.96 112.87 ± 18.78b 5.07 ± 0.84
Season
Spring 203.81 ± 77.66b 24.21 ± 9.56 0.93 ± 0.30b
Summer 216.89 ± 77.70ab 25.52 ± 9.56 0.98 ± 0.30ab
Fall 314.57 ± 72.34a 37.04 ± 9.02 1.41 ± 0.30a
Winter 283.00 ± 76.71ab 33.39 ± 9.50 1.26 ± 0.32ab

(i.e. sites with high basal area and low thinning intensity with a mean basal area of 19.03 m2
ha−1), although these sites also had the greatest variability in litter biomass (Table 3). The
carbon content of litter was highest in the H50 treatment (the sites with highest residual
basal area) and was significantly lower in the two treatments with lower residual basal area
(L50 and L75) (Table 3; Tukey pair-wise comparison, p < .05). For nitrogen content, there
were no significant differences across treatments.
Across all sites, we found significantly less litter biomass falling in the spring (203.81 kg
ha−1) compared to the fall season (314.57 kg ha−1), while summer and winter litter falls were
not significantly different from other seasons because of the high variance across sites in
these two seasons (Table 3; Tukey pair-wise comparison, p < .05).

Cluster analysis of litterfall characteristics by site

In the hierarchical cluster analysis, two distinct groups emerged with two sites as outliers
(Figure 3). Although the sites did not clearly group based on thinning treatment, there were
patterns in stand characteristics among clusters. The orange cluster, which also had lower
biomass, carbon and nitrogen, also showed high basal area, with a mean of 17.2 m2 ha−1 and
a high mean tree density of 645 trees ha−1. The green cluster, which had the highest mean
biomass and carbon, and high nitrogen, showed lower basal area and lower tree density
(mean BA = 14.1 m2 ha−1 and mean density = 506 trees ha−1). Finally, the outlier to the
8 Y. PÉREZ-ALAVEZ ET AL.

Figure 3. Hierarchical cluster analysis of sites based on litter biomass, nutrient content, and site
characteristics. Sites are labeled by number and treatment, with 50 or 75 for thinning intensity
(the percent of BA removed) and h for high residual BA (˃13 m2 ha−1) and l for low (≤13 m2 ha−1).
Two distinct groups are highlighted with a green oval and an orange rectangle.

green cluster showed the second highest biomass and carbon, and the highest nitrogen
levels, and also had the lowest BA (10.7 m2 ha−1) and density (325 trees ha−1). The outlier to
all clusters, site 3, had the lowest biomass, carbon and nitrogen of the clusters, and also had
very high basal area and high tree density compared to other sites (BA = 22.4 m2 ha−1 and
density = 725 trees ha−1). This analysis therefore shows an association between lower BA
and density and higher leaflitter biomass and nutrient content, with the two outliers as sites
that break with this pattern.

Regression modeling of litter characteristics

Linear regression models showed no significant relationship between litter biomass or
nutrient concentrations and tree density, but there were significant relationships at the
p < .1 level between litter characteristics and post-thinning basal area. Litter biomass was
positively correlated with basal area (Figure 4; linear regression, adj R2 = 0.20, p = .08), and
litter carbon concentration was also positively correlated with basal area (linear regression,
adj R2 = 0.26, p = .051).

Discussion
This study is one of the first to examine litterfall in managed natural pine forest in Oaxaca,
Mexico. We found that thinning intensity had a significant impact on litter carbon, with
significantly higher carbon in the sites under the least intensive thinning treatments (50 H
Figure 4. Linear regressions of (a) litter biomass by site basal area and (b) litter carbon content by site basal area, with regression equations, the adjusted R2 values
and the p-values.
JOURNAL OF SUSTAINABLE FORESTRY
9
10 Y. PÉREZ-ALAVEZ ET AL.

sites), and that residual basal area was positively correlated with higher litter biomass in the
linear regression and was associated with higher litter biomass in the hierarchical cluster
analyses. Furthermore, we found distinct seasonal patterns in litterfall, with peaks at the
onset of the dry season in October-December and at the end of the dry season in March.
Unfortunately, little research exists on litterfall in similar ecosystems in southeastern
Mexico, and we found no other studies examining the impacts of thinning on litter in this
region or forest type. Other litterfall research in Mexico has focused on tropical dry forest
(Lawrence, 2005; Martinez-Yrizar & Sarukhan, 1990; Morffi-Mestre et al., 2020; Saynes
et al., 2005), tropical semi-evergreen forest (Aryal, 2015), cloud forest (Williams-Linera
et al., 1996), deciduous Acacia plantations (Ngangyo-Heya et al., 2017), or even mangrove
systems (Aké-Castillo et al., 2006; Arreola-Lizárraga et al., 2004; González-Rodríguez et al.,
2019), but less research examines this type of temperate pine-oak forest. Only González-
Rodríguez et al. (2019), González-Rodríguez et al. (2018), p. 2011) and Pérez-Suarez et al.
(2009) examine litter in pine and pine-oak forests in Mexico, although in northeastern and
northwestern Mexico, respectively, and we could find no published literature on this forest
type in Oaxaca state in southwestern Mexico. Our study therefore contributes to a better
understanding of nutrient cycling dynamics in this understudied but economically impor­
tant forest type in Oaxaca, Mexico.

Impact of thinning on litterfall

Although we did not consistently find significant differences in litter properties across all
treatments in our analyses, we did find evidence that thinning has an impact on litter
quantities and that sites with higher residual basal area produce more litter in this stand.
There were significant differences in carbon content between thinning treatments, with
more carbon deposited in litter in the least intense of the thinning treatments (H50).
Similarly, the cluster analysis grouped sites together such that lower basal area and lower
tree densities were associated with higher litter biomass, carbon and nitrogen. Furthermore,
there were positive correlations between annual litter biomass, litter carbon content and site
basal area (which ranged from 8.96 to 24.5 m2 ha−1) at the p < .1 significance level. Other
studies of the effects of thinning on litterfall have found conflicting results around the
world, with thinning leading to changes in litterfall in some cases and no differences in
others, implying that the impacts of silvicultural management on litter dynamics are
complicated by factors such as stand age, species, climate patterns, and the management
intervention used (Çömez et al., 2019; Jiménez & Navarro, 2016; Novák et al., 2020; Segura
et al., 2017).
One possible explanation for the lack of a clear difference in litter biomass between
thinning treatments in this study is that the impact of thinning on litterfall had diminished
in the intervening two-three years between the treatment and our study. The time-frame for
any impacts of thinning on litterfall are unclear, as one study of pine stands in Italy found
that the short term impacts of thinning disappeared after the first year (Lagomarsino et al.,
2020), while other studies in Spain found that thinning effects on litterfall remained after
five to eight years (Jiménez & Navarro, 2016; Lado-Monserrat et al., 2015; Navarro et al.,
2013). Another study in Spain did find that unthinned stands produced more litter
compared to thinned stands, but that the effect of thinning in decreasing litterfall had
disappeared after 5 years (Roig et al., 2005).
 JOURNAL OF SUSTAINABLE FORESTRY 11

The type of pine stand also seems to determine whether thinning impacts litterfall, both
in terms of species composition, site characteristics, and stand maturity. For example, in the
Czech Republic the impact of thinning on litterfall was much less dramatic in stands
dominated by pine alone compared to those that were mixed pine-Douglas fir (Novák
et al., 2020). In Spain, the effects of thinning on litterfall were site dependent such that one
site showed significant differences between thinning intensities, while another showed no
differences (Blanco et al., 2006). In Turkey, thinning treatments had an impact on litterfall
in mature stands, but not in overmature stands (Çömez et al., 2019).
The relationship between litterfall and basal area may also be exacerbated as stands
mature. Thus while Kim (2016) found that leaf litter was positively correlated with basal
area, their natural stands in Korea had basal area stocking rate way above those of this
P. pseudostrobus stand in Mexico. Our study examined quite a young stand, so potentially
the impacts of differing thinning regimes here were less clearly visible than they would be at
a more mature stage. Furthermore, many of these other studies occurred in plantations
rather than naturally regenerated stands, which may introduce other differences depending
on the management and stocking levels. While it would certainly be ideal to compare our
findings to other natural forests under silvicultural management, there is little other
research publicly available on litterfall fluxes in natural pine stands.
Although thinning may tend to decrease the amount of litterfall due to the reduction in
the number of trees producing litter, some of these impacts may be ameliorated by resulting
growth in the foliar biomass of residual trees. Thus Rodríguez-Ortiz et al. (2011) found that
different thinning intensities significantly increased foliar biomass in a P. patula stand in
Oaxaca, Mexico, with greatest increases in foliar biomass associated with the higher thin­
ning intensity of 68% of density. According to models created in similar pine stands in the
region, the thinning treatments applied here of 50% and 75% should lead to similar
increases in leaf area and foliar biomass (Rodríguez-Ortiz et al., 2012), which may be
causing the lack of significant difference in litter biomass between stands with more basal
area removed and those with more basal area left residually. According to studies of biomass
growth in México, young stands between 10 and 20 years in age incorporate carbon and
biomass at higher rates, so the resulting growth after thinning in this young stand of
12 years may be quite high, thereby complicating the impacts of thinning on litter (Casiano-
Domínguez et al., 2018).
The mixed results in nutrient fluxes between thinning treatments (i.e. the significant
difference in litter C between treatments but not in litter N) demonstrate that nutrient
patterns in litterfall are not always straightforward or predictable. Although Kim (2016)
found negative relationships between basal area and C and K fluxes in naturally regenerated
stands, there were positive relationships between basal area and other nutrients. Other
research has also found significant relationships between thinning treatments and some
nutrient fluxes but not for others in litter in pine plantation systems (Segura et al., 2017)
or increases in some nutrients and decreases in others (Bueis et al., 2018). Lastly, studies show
that even if thinning can impact litter quantities, these treatments do not seem to impact soil
carbon (Ruiz-Peinado et al., 2013) or soil nutrient concentrations (Bravo-Oviedo et al., 2013).
Finally, this study had a few limitations that may have impacted our conclusions. First
and foremost, we lacked unthinned control plots to compare to our different thinning
intensities. Because this study occurred on actively managed community forest, the study
design was constrained by the existing forest management, which had required thinning on
12 Y. PÉREZ-ALAVEZ ET AL.

all parts of this young stand. However, thinning is a common management activity in this
and other forested areas, so understanding the impacts of thinning intensity is in many ways
a more practical and realistic concern for actively managed stands. Balancing timber
production with ecosystem services and carbon sequestration will be increasingly important
for the adoption of effective conservation and climate change mitigation measures, and so
studies that contribute to understanding the impacts of silvicultural management on carbon
and nutrient dynamics will be essential for models and analyses that assess the relative
production, ecosystem, and climate impacts of different management systems (Bonan, 2008;
Canadell & Raupach, 2008).

Litterfall biomass and nutrient fluxes

The accumulated litter biomass of 1059.27 kg ha−1yr−1 in a thinned 10-year-old
P. pseudostrobus stand found here in Oaxaca was comparable to that reported in other
pine stands in seasonally dry forests. These values are particularly aligned with studies of
Aleppo pine (P. halepensis) stands thinned at a 75% treatment in the Mediterranean region,
which have found a range in total annual litterfall from 950 kg ha−1 yr−1 (Navarro et al.,
2013) to 1300 kg ha−1yr−1 (Jiménez & Navarro, 2016). Other research in pine forests in
Europe have found comparable or lower amounts of litterfall, such as a range from 950 to
2280 kg ha−1 of dry matter for P. halepensis (Bueis et al., 2018), a mean of 635 kg ha−1 for
P. halepensis and a mean of 836 kg ha−1 for P. sylvestris in stands in Spain (Bueis et al., 2017).
Finally, while there appears to be a great range in litterfall quantities depending on the
maturity of the stand, the quantities reported here for a 12-year-old stand align well with
those reported in young Scots pine stands in Turkey (1389 kg ha−1 yr−1) but are much lower
than those reported for mature stands (4488 kg ha−1 yr−1) (Çömez et al., 2019).
The accumulated annual carbon and nitrogen contributions from litterfall in this study
of 125.32 kg ha−1 of carbon and 4.76 kg ha−1 of nitrogen were comparable to reported values
in some studies but lower than those in other forests, likely because of the age, stocking, and
species composition of the stand. The annual carbon flux in litter in this P. pseudostrobus
stand was higher than that reported in a P. pinaster stand in Spain, where annual
C contribution from litter was 49.9 kg ha−1 (Bravo-Oviedo et al., 2013). On the other
hand, annual carbon and nitrogen were lower compared to annual means in litter in an
Aleppo pine stand in Spain of 536.5 kg C ha−1yr−1 and 6.5 kg N ha1yr−1 (Segura et al., 2017),
and lower compared to mature natural P. densiflora stands in Korea with much higher
stocking rates where N ranged 13.84 to 20.68 kg ha1yr−1 (Kim, 2016). Nitrogen contribu­
tions from litter also appear to be lower in this stand compared to two mixed pine-oak
stands in northern México studied by González-Rodríguez et al. (2011) which had 18.3 kg
N ha−1 yr−1 and González-Rodríguez et al. (2019) with 27 kg N ha−1 yr−1 from litter.
However, in these cases the addition of deciduous leaf litter likely increases both overall
quantities of litter as well as nutrient concentrations within.

Seasonality of litterfall
In this pine stand in southern México, we found the greatest litterfall in fall and winter from
October – December, with a secondary peak in March. These results align well with studies
of litterfall in northeastern México, where mixed pine-oak forest showed similar seasonal
 JOURNAL OF SUSTAINABLE FORESTRY 13

peaks in March and November (González-Rodríguez et al., 2011), and pine forest showed
peaks in April/ May and December (González-Rodríguez et al., 2019, 2018). In contrast,
a study of P. cembroides in northwestern Mexico found a peak in June, so there may be site-
specific and species-specific differences even within the same general climate type (Pérez-
Suárez et al., 2009). For example, in a five year study of deciduous tropical dry forest in
Jalisco (another state on the Pacific coast of Mexico), seasonal litterfall peaked near the
onset of the dry season at one more xeric site and but peaked in the middle of the dry season
at another more mesic site due to differences in underlying soil moisture availability
(Martinez-Yrizar & Sarukhan, 1990)
Although México and the Mediterranean demonstrate key differences in the timing of
dry seasons, global analysis shows that seasonal patterns in litterfall tend to coincide with
dry season (Zhang et al., 2014). In Europe and the Mediterranean, seasonal peaks in pine
litterfall also tends to occur in the dry season, although there has been some variation across
species and studies. For example, P. pinaster stands showed the greatest peak from August
to October (Roig et al., 2005), P. sylvestris stands have shown peaks in September/October
(Blanco et al., 2006) and in July-September (Bueis et al., 2017), and P. halepensis stands have
shown primary peaks in August-September (Bueis et al., 2017; Navarro et al., 2013). In the
Mediterranean, seasonal droughts tend to occur during the summer, whereas in south­
western México, the dry season begins in the fall (usually October or November) and lasts
through the end of March, although it can extend to May or June in some years
(CONAGUA, 2020). Thus the two peaks of October-December of and March may represent
two periods of drought stress which led to greater needle fall in this stand.
Another possible explanation for the October-December peak, however, is that the
higher litterfall in autumn in this pine stand was triggered by the onset of the colder winter
temperatures, and that the litterfall was following a more temperate seasonal pattern.
A limitation to the study is the relatively short time period of monitoring (one year).
Monitoring litterfall for a greater time period and across a greater range of locations in
southwestern Mexico would help elucidate linkages between litterfall, drought, and tem­
perature. It is possible that seasonal patterns may shift from year to year, and so further
study of litterfall cycles would certainly be valuable to confirm the relationship between
litterfall and dry and wet seasons. Additionally, as the impacts of thinning on litter likely
shift with time since the thinning operation (Roig et al., 2005), a longer study could provide
a stronger assessment of the lasting impacts of management on nutrient cycling dynamics.

Conclusions
This study found clear seasonal patterns in litterfall in pine forests in southwestern Mexico,
with the highest peak in the dry season in autumn/early winter when 58.22% of all litter
biomass was deposited. However, the impacts of thinning treatment on litterfall were less
straightforward, with significant impacts of thinning intensity on carbon content and
correlations between residual basal area with litter biomass, but no significant impacts on
nitrogen. We also found that the biomass and nutrient contributions from litter were
comparable to those reported in similar seasonally dry pine stands in the Mediterranean.
Litter deposition is impacted by many factors, including stand age (Çömez et al., 2019),
elevation and successional status (Zhou et al., 2007), and climatic factors (Berg &
Meentemeyer, 2001), and it seems likely that the young age of this stand, as well as the
14 Y. PÉREZ-ALAVEZ ET AL.

increased growth triggered by release from competition, may complicate the impacts of
basal area removal on litter production.
Finally, this is one of the first studies to examine seasonal litter dynamics in this ecologi­
cally and economically important forest type in México. While there were some limitations to
this study, notably the lack of an unthinned control and the relatively short time frame, we
present a starting point for further research on the impacts of silvicultural treatments on
carbon and nutrient cycling patterns in this system. Forests are a large and persistent carbon
sink, and litter plays a small but not insignificant role in the forest carbon and nutrient cycle
(Pan et al., 2011). The best way to manage forests for climate mitigation, and the potential
role of timber production within this framework remains debated (Bellassen & Luyssaert,
2014; Creutzburg et al., 2017), and this research contributes to our understanding of the
impacts of silvicultural thinning regimes on carbon dynamics. Furthermore, accurate assess­
ments of carbon cycling under different silvicultural treatments will be useful for commu­
nities interested in taking part in future carbon markets. Lastly, climate change has the
potential to impact the timing and severity of the dry season in Mexico, and so understanding
the seasonality of nutrient cycling provides an important baseline for future monitoring.

Acknowledgments
We would like to thank the commissary of communal possessions and forestry technical services
(Bienes Comunales y Servicios Técnicos Forestales) of the San Pedro el Alto community for use of
their communal forest for this research. Thank you to Eng. Moisés Raúl Hernández C. of San Pedro el
Alto for his support and assistance as well.

Disclosure statement
The authors have no conflicts of interest to disclose regarding this research.

ORCID
Wenceslao Santiago-García http://orcid.org/0000-0003-1958-1696
José Raymundo Enríquez-del Valle http://orcid.org/0000-0002-7700-3790
Meredith P Martin http://orcid.org/0000-0001-5583-3390

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