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Questão 1 (0.5 pontos)
Explique com suas próprias palavras (e podemos fazer esquemas) a diferença entre 1) variáveis
explanatórias (independentes) e variáveis resposta (dependentes) e 2) variáveis teóricas e
variáveis operacionais.
Questão 2 (0.5 pontos)
Sobre as variáveis operacionais, explique, com suas próprias palavras, a diferenças entre variáveis
quantitativas, variáveis categóricas (qualitativas) e variáveis semi-quantitativas (ordinais).

Questão 3 (1.0 pontos)

Junto a esta lista você recebeu em formato PDF um artigo científico (Moreno et al. 2014 Area and edge
effects on leaf-litter decomposition in a fragmented subtropical dry forest. Acta Oecologica 60: 26-29).
Em relação a este artigo, responda:
a) Quais eram as perguntas que os autores do artigo buscaram responder?
b) Quais eram as variáveis explanatórias (teóricas e operacionais) deste estudo? Elas eram
quantitativas, qualitativas ou semi-quantitativas?
c) Quais eram as variáveis resposta (teóricas e operacionais) deste estudo? Elas eram quantitativas,
qualitativas ou semi-quantitativas?

Questão 4 (0.5 pontos)

Ainda em relação ao artigo, a que conclusões os autores chegaram sobre as relações entre as variáveis?
 Acta Oecologica 60 (2014) 26e29

Contents lists available at ScienceDirect

Acta Oecologica
journal homepage: www.elsevier.com/locate/actoec

Short communication

Area and edge effects on leaf-litter decomposition in a fragmented

subtropical dry forest
M.L. Moreno a, b, *, M.L. Bernaschini a, b, N. Pe

rez-Harguindeguy b, G. Valladares a, b
a
Centro de Investigaciones Entomolo
gicas de Co

rdoba, F.C.E.F.yN., Universidad Nacional de Co

rdoba, Av. V

rdoba,
elez Sarsfield 1611 (X5016GCA), Co
Argentina
b
Instituto Multidisciplinario de Biología Vegetal (CONICET-Universidad Nacional de Co
rdoba), Av. V

elez Sarsfield 1611-(X5016GCA), Facultad de Ciencias
Exactas Físicas y Naturales, Universidad Nacional de Co
rdoba, Argentina

a r t i c l e i n f o a b s t r a c t

Article history: South American subtropical dry forests are highly threatened by fragmentation. Despite considerable
Received 2 December 2013 research efforts aimed at predicting ecosystem alterations due to this driver of global change, we still
Accepted 24 July 2014 need to deal with general principles to improve our ability to predict the impact of fragmentation. Our
Available online 30 August 2014
work is one of the few studies that analyse the relationship between forest fragmentation and decom-
position. In 12 remnants of Chaco Serrano forest in Central Argentina we tested if decomposition rates of
Keywords:
a common leaf-litter substrate varied with fragment size and between the forest edge and interior.
Area loss
Decomposition declined with fragment size, with no significant effects of location (edge/interior) or
Forest edge
Habitat fragmentation
interaction between the two components of fragmentation. Our results suggest that in situ conditions for
Discontinuous canopy decomposition may change as a consequence of forest fragmentation, specifically as a result fragment
Chaco Serrano size. This may lead to impaired nutrient recycling in smaller forest remnants.
© 2014 Elsevier Masson SAS. All rights reserved.

1. Introduction reach a generalization, Fahrig (2003) showed that habitat loss had
larger and consistently negative effects on biodiversity while
South American tropical and subtropical dry forests are dis- different components of habitat fragmentation per se (e.g. isolation,
appearing at an alarming rate, with 80% deforestation of the orig- edge effects, matrix quality) had weaker effects, which could be
inal area being recorded in the last decade (Aide et al., 2013). either positive or negative. Didham et al. (2012) proposed that in
Deforestation involves habitat loss and fragmentation leading to spite of the magnitude of each effect on biodiversity, they all
the transformation of a forested area to a number of smaller forest operate through chains of direct and indirect causal links which
remnants embedded in a matrix with different land use, usually should be addressed by fragmentation studies. How the different
agricultural (Fahrig, 2003). Although ongoing habitat loss and components of fragmentation alter ecosystem processes has not
fragmentation are recognized as primary drivers of contemporary been studied extensively particularly in South American ecosys-
extinctions (Brook et al., 2008), we still need to deal with general tems (Grez and Galetto, 2011).
principles to improve our ability to predict the impacts of processes As forest remnants become smaller, they become increasingly
involved in land use change. In order to achieve this goal, infor- dominated by edge habitat (Ries et al., 2004). Remnants edges can
mation from field studies representing different ecosystems is a present particular microclimatic characteristics such as reduced
fundamental requirement. humidity, greater temperature variability and increased solar ra-
There has been a long debate on the distinction between habitat diation in comparison with the forest interior (Laurance, 2004).
loss and habitat fragmentation per se (Banks-Leite et al., 2010; Consequently, smaller forest remnants, with a high proportion of
Ewers and Didham, 2007; Haila, 2002). In a valuable attempt to edge habitat, may be, on average, hotter and drier compared to
larger ones. However, microclimate changes between the edge and
interior could be more complex (Davies-Colley et al., 2000;
Gehlhausen et al., 2000), with even opposite trends in microcli-
* Corresponding author. Centro de Investigaciones Entomolo
gicas de Co

rdoba,
Instituto Multidisciplinario de Biología Vegetal, Edificio de Investigaciones Bio-
matic factors being recorded (Wright et al., 2010). In spite of the

gicas y Tecnolo
lo
gicas, Av. Ve

lez Sarsfield 1611-(X5016GCA) Co
rdoba, Argentina. intrinsic relationship between remnant area and edge/interior
Tel.: þ54 351 6653351.
E-mail address: [email protected] (M.L. Moreno).

http://dx.doi.org/10.1016/j.actao.2014.07.002
1146-609X/© 2014 Elsevier Masson SAS. All rights reserved.
 M.L. Moreno et al. / Acta Oecologica 60 (2014) 26e29 27

location, most studies focus either on the ecological responses to between 31100 e31300 S and 64 000 e64 300 W, at 500e600 m
habitat loss or to edge effects. a.s.l., twelve forest fragments which had been isolated for at least
Leaf-litter decomposition is a fundamental process that con- 30 years, ranging in size from 0.57 ha to over 1000 ha were selected.
tributes to carbon and nutrient cycling and hence to ecosystem
productivity (Ha €ttenschwiler et al., 2005; Zhang and Zak, 1995). 2.2. Litter bag experiment
This process is hierarchically controlled by climatic conditions (e.g.
temperature and humidity Coûteaux et al., 1995), leaf quality We collected freshly senesced leaf material of two abundant
(Cornelissen et al., 1999) and soil organisms (Gonzalez and native species (Celtis ehrenbergiana (Klotzsch) Liebm. and Croton
Seastedt, 2001). The relationship between habitat fragmentation lachnostachyus Baill.) during the peak of leaf fall (MayeAugust
and decomposition has been addressed in very few studies and the 2009), within the largest forest remnants (>1000 ha). We deter-
results reported have been inconclusive. For example, decomposi- mined initial leaf-litter quality of those two common substrates
tion was slower at the edge in comparison with the interior of (see Table 1). Samples of the two species were oven-dried at 50  C
temperate mixed deciduous forests (Riutta et al., 2012), whereas in during 48 h and ground for chemical analysis (see Table 1). We
Andean cloud forests (Romero-Torres and Varela Ramírez, 2011) determined total carbon (C) as 50% of ash-free biomass
and Australian dry forests (Hastwell and Morris, 2013) the effects (Schlesinger, 1977; McClaugherty et al., 1985; Gallardo and Merino,
on decomposition depended on edge orientation. Similarly variable

1993), while total nitrogen (N) (ONeill and Webb, 1970), lignin (L),
responses to edge conditions have been reported for tropical sys- cellulose and hemicellulose (Goering and Van Soest (1970) were
tems (Didham, 1998; Noble, 2013; Rubinstein and Vasconcelos, measured with an Autoanalyser (RFA 300-Alpken, Wilsonville, OR,
2005). With regard to fragment size, different studies have found USA) at the Chemical Analyses Laboratory of INTA Bariloche. The
either a decline in decomposition with forest area (Didham, 1998; litter was air-dried and thoroughly mixed to form one common
Wardle, 1997) or no effects (Hastwell and Morris, 2013; Noble, pool with equal amount of each species.
2013; Rubinstein and Vasconcelos, 2005). We know of no study Litter bags (24  21 cm; n ¼ 72) were constructed with 0.3 mm
simultaneously addressing the effects of edge/interior location and nylon mesh on the bottom side, and 1 mm nylon mesh on the upper
fragment area on decomposition in subtropical dry forests, except side. The thinner mesh prevents leaf-litter loss from the bottom
for the work by Hastwell and Morris (2013) in which edge effects side, whereas the coarser mesh allowed the access of micro and
were thoroughly explored but area variations were represented by mesofauna through the top side of the bag. The litter-bags were
four forest fragments. filled with 6 ± 0.02 g of the common substrate pool. In order to
In this context, we experimentally analysed leaf-litter decom- estimate true dry mass before litter-bag burial we first calculated
position along an area gradient in a fragmented Chaco Serrano forest air-dried water content. We air-dried five sub-samples of the leaf-
in central Argentina. This system is part of the most extensive and litter substrate and oven-dried them at 50  C for 48 h. Water con-
highly threatened dry forest in South America (Zak et al., 2004). We tent value (%) was deducted from air-dry mass in all samples.
were interested in whether decomposition can be affected by forest In December 2009 six litter-bags were places on the soil surface
area and edge effects. By using a common substrate (from native of each of the 12 forest fragments, in two locations: edge (within
plants) we avoided effects mediated by changes in litter quality, thus 2 m from the end of the tree line) and interior (about 30 m from the
focussing on effects of in situ conditions for decomposition. end of the tree line). Each litter-bag was positioned with the thin
mesh side in contact with soil, and the wide mesh side was covered
2. Materials and methods with some of the natural litter layer of each fragment to simulate
natural decomposition conditions. We retrieved all litter-bags after
2.1. Study area 120 days of incubation and immediately transported them in paper
bags to the laboratory, where they were stored at 4  C until
The study was conducted in the Chaco Serrano District, in cen- processing. Samples were then defrosted, cleaned by removing soil
tral Argentina, a seasonally dry forest with a mean temperature of material, and oven-dried at 60  C for 48 h for dry mass determi-
17.5  C and an average annual rainfall of 750 mm mostly concen- nation. To correct for soil contamination, the remaining ash free dry
trated in the warm season (OctobereApril) (Fig. 1). The native mass (AFDM) for each litter-bag was measured by incineration at
vegetation, currently reduced to isolated remnants embedded in a 550  C for 4 h (Harmon and Lajtha, 1999). Decomposition rate was
predominantly agricultural matrix, includes a tree layer (8e15 m estimated as the difference between initial litter dry weight and
high), shrubs (1.5e3 m), herbs and grasses (0e1 m) and many vines litter dry weight after incubation, without ash.
and epiphytic bromeliads (Cagnolo et al., 2006). In an area located
2.3. Data analysis

To evaluate the effects of habitat fragmentation on leaf-litter

decomposition we developed a linear mixed model (Pinheiro and
Bates, 2000) with decomposition rate (% dry weight mass loss) as
response variable. Fragment size (ha) and location (edge/interior)
were treated as fixed effects. Site (fragment identity) was included

Table 1
Initial litter quality of two common plant species from Chaco Serrano forest.

Nitrogen Carbon Lignin Cellulose Hemicellulose

(%) (%) (%) (%) (%)

Celtis ehrenbergiana 1.76 71.40 5.1 14.9 0.1

(Klotzsch)
Fig. 1. Climodiagrams representing mean monthly temperatures and precipitation for
Croton lachnostachyus 1.49 42.36 6.3 23.7 1.6
the study site, taken from closest meteorological station (6 km from the fragments,
(Baill)
Servicio Metereolo
gico Nacional Argentino) during the period of the experiment.
28 M.L. Moreno et al. / Acta Oecologica 60 (2014) 26e29

as random factor to account for the non-independence of edge and

interior in the same fragment. Location (interior/edge) was also
incorporated as nested factor within site to contemplate the co-
dependence of the three litter-bags from each location. We also
evaluated interactions between fixed effects. Non-significant in-
teractions were removed in order to obtain the most parsimonious
model. Statistical analyses were performed using R 2.11.0 software
(R Development Core Team, 2010). After checking data distribution,
values were arcsine (percentage dry mass loss) or log (fragment
size) transformed in order to achieve normal distribution.

3. Results

After 120 days of incubation (December 2009eApril 2010) in the Fig. 2. Leaf-litter dry weight mass loss (%) after 120 days of incubation at edge and
wet and warm season of Chaco Serrano (see Fig. 1), the decompo- interior locations in forest remnants of Chaco Serrano.
sition of the common substrate varied between 50 and 85% mass
loss (mean ¼ 72.80 ± 1.41%). As forest fragment size decreased,
decomposition rates also declined (Table 2, Fig. 2). Decomposition
rates did not differ between the forest edge and the interior litter quality (via changes in plant species abundance and identity,
(Table 2). The effects of forest area and edge/interior location on Noble, 2013) or the decomposer biota (Rantalainen et al., 2005). In
decomposition rates were independent from each other since the our experiment, litter quality was kept constant by using the same
interaction between these factors was not significant (F ¼ 0.46; substrate in all fragments, so it is not a change in litter quality that is
p ¼ 0.51). causing slower decomposition in smaller fragments (Didham,
1998). Instead, in situ conditions and particularly the role of the
4. Discussion decomposer biota in decomposition remain to be investigated. A
reduction in fragment area has been linked to impoverished
In one of the first studies of leaf-litter decomposition simulta- detritivorous fauna in rainforest systems (leaf litter beetles:
neously testing area and edge effects in a fragmented subtropical Didham et al., 1998; termites: Davies, 2002; leaf-litter ants:
dry forest, we have shown decomposition rates of a common Carvalho and Vasconcelos, 1999), with exclusion of soil in-
substrate declining with forest fragment size but no effect, or vertebrates resulting in decreased leaf litter decomposition
interaction, of location within fragment. Changes in fragment area (Bradford et al., 2002; Meyer et al., 2011; Powers et al., 2009). A
could affect environmental conditions for decomposition via edge recent study found that slow decomposition of common substrates
effects: as fragments become smaller, conditions in the interior in forest patches was linked to changes in macroinvertebrate
approach those in the drier and hotter edge habitats. In such a communities (Cuke, 2012).
scenario, decomposition should be expected to decrease at the In our study we did not detect any edge effects on litter
edges and in smaller remnants, with integration between edge and decomposition. Edge effects on leaf-litter decomposition appear to
interior in the latter, where edge conditions become prevalent. be highly variable in different systems. For example, in tropical
However, after 120 days of incubation, these hypotheses were not systems decomposition was either highest at forest edges or did not
supported by our results, since decomposition rates did not differ differ with position (Didham, 1998; Rubinstein and Vasconcelos,
between forest edge and interior habitats, nor did edge interact 2005). The opposite pattern was found in temperate forest (Riutta
with forest area. Conversely, Hastwell and Morris (2013) found no et al., 2012). Cardinal orientation of edges may also influence
effect of area on leaf-litter decomposition in an Australian forest, decomposition rates of litter (Romero-Torres and Varela Ramírez,
whereas decomposition rates decreased near edges depending on 2011; Hastwell and Morris, 2013). Such variability could be
edge orientation. related to the complexity of interactions between microenviron-
Another possible mechanism for slower decomposition rates in ment and some characteristics of edge habitat. For example,
smaller fragments could be related to species loss, affecting either increased temperature in forest edges may either enhance or slow
decomposition depending on moisture content, which may in turn
be modulated by edge orientation, altitudinal variations, type of
Table 2 matrix, among other factors (e.g. Gehlhausen et al., 2000; Romero-
Details of linear mixed models examining the relationship between edge/interior
Torres and Varela Ramírez, 2011). To make the picture more com-
localization, fragment area (log) and dry weight mass loss (arcsine %) after 120 days
of incubation in remnants of Chaco Serrano. Location (edge/interior) and fragment
plex, most studies on microclimatic changes between edge and
area were modelled as fixed effects whereas site (fragment identity) and location interior habitats have focused on tropical systems (Laurance, 2004),
(interior/edge) nested within site, as random effects. SE (standard error), DF (de- whereas recent evidence has shown fragment interior being
grees of freedom). warmer and drier than edges in forests with discontinuous canopy
Term Estimates ± SE DF F p StDev Residuals cover, such as in Chaco Serrano (Wright et al., 2010). Finally,
Intercept 1.01 ± 0.01 1, 48 12876.32 <0.0001
detection of edge-related changes depends on the distance used to
Location -0.02 ± 0.02 1, 11 1.48 0.24 define edge and interior habitats. In some systems, edge microcli-
(Edge/ matic changes can penetrate more than 40 m into the interior of a
Interior) forest (Kapos, 1989; Davies-Colley et al., 2000). The distance be-
Fragment 0.04 ± 0.008 1, 10 18.09 0.002
tween edge and interior samples in our study (30 m) might be
area
Random deemed insufficient to perceive edge effects on leaf litter decom-
effects position related to microclimate. Nonetheless, a similar design
Site identity 2.86E-06 allowed detection of edge-related differences in herbivory and
Site identity 4.18E-07 0.07 parasitism rates in forest remnants in Chaco Serrano (Valladares
(Location)
et al., 2006).
 M.L. Moreno et al. / Acta Oecologica 60 (2014) 26e29 29

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