Evaluation of native woody species performance related to environmental

conditions and plant functional traits in rehabilitation of degraded areas in

the northeast of Patagonia, Argentina.
J M Zeberio a,b, C A Pérez c.
a
Universidad Nacional de Río Negro, Sede Atlántica. Centro de Estudios Ambientales desde la Norpatagonia (CEANPa). Viedma,
Río Negro.

b
National Council of Scientific and Technical Research (CONICET).

c
Laboratorio de Investigación de Sistemas Ecológicos y Ambientales (LISEA). Universidad Nacional de La Plata. La Plata, Buenos
Aires.

Abstract. Degradation processes affect vast area of arid and semiarid lands around the world and cause
damages in environment and health people. Degradation processes are impulsed for human produtive
activities that cause direct and indirect effects on natural resources as species extintion to regional or predial
scale, reduction and elimination of vegetation cover, soil erosion, among others. In front of this, ecological
rehabilitation is an important tool to recover key aspects from degraded ecosystem. To develop rehabilitation
trials is desirable use native plant species of local procedence with characteristics that allow them high
survival and growth rates. The aim of this work was to assess the survival and growth of native woody species
in degraded areas of the northeast of Patagonia and relate them to plant functional traits and environmental
variables. We observed high early and late survival rates and growth by Prosopis flexuosa and Schinus johnstonii,
and low values by Condalia microphylla and Geoffroea decorticans. Early survival rates are positevely associated with
Specific Leaf Area (SLA) and precipitation, and negatively associated with wood density and maximum mean
temperature of January and minimum mean temperature of July. Late survival rates were positevely associated
with SLA and soil organic matter, and negatively associated with plant height and precipitations.
Temperatures had a positive effect on late survival rates once plants overcame the critical threshold of the
first summer after they were transplanted to the field. Prosopis flexuosa and S. johnstonii were the most
successful species of our study. This could bedue to their functional traits that allow this species to
acclimatize to local environments charecteristics. It is desirable to continue the researchs about C. microphylla
and G. decorticans to know aspects related to productive conditions, rustification processes, autoecology and
potential usefull in ecological rehabilitation trials.

Keywords: Arid lands; Condalia microphylla; Geoffroea decorticans; Prosopis flexuosa; Schinus johnstonii; Survival rates;
heigh Growth, basal diameter growth.

1 Introduction

Degradation processes affect vast areas of arid and semiarid lands around the world, which are
habited by 2.5 billions of people (UNCCD, 2011). Most of them are in developing countries, with
areas that present several degradation symptoms, and impoverishment health of people and of
ecosystems (James et al., 2013).

Usually, the causes of degradation of ecosystems are related to human activities and are evidenced
by losses of vegetation cover, soil erosion and soil fertility, which diminishing land productivity
(Cortina et al., 2011; Vallejo et al., 2009; Abraham et al., 2009). In this scenario, ecological
rehabilitation is a key strategy to recover degraded ecosystems (Aronson et al., 1993; Delgado-
Baquerizo et al., 2013; Maestre et al., 2001).

The ecological rehabilitation performs tasks on natural system to return some of its lost structural
and functional attributes. To achieve this objective, disturbance factors should be removed, and
actions should be taken to increase vegetation cover (Aronson et al., 1993, Hobbs and Kramer,
2008). Different techniques are commonly applied to attain the last propose, such as sowing seeds of
native or exotic species in bare soils, transplanting plants to the field and/or applying soil
amendments to improve physical and chemical soil properties (Dalmasso, 2010; Landi and Renison,
2010; Mcadoo et al., 2016; Meli et al., 2013; Soteras et al., 2014; Torres et al., 2015; Delgado-
Baquerizo et al., 2013; Milgrom, 2008; Neri and Sánchez, 2010; Novak and Prach, 2003; Gasch et
al., 2014; de Villalobos et al., 2001, 2005; Maestre et al., 2001; Peláez et al., 1996; Pérez et al.,
2019; Plaza Behr et al., 2016). Actively plantation of species produced in plant nursery is one of the
most frequently used techniques in rehabilitation trials (Perez et al., 2019). In addition, it is desirable
to initiate ecological rehabilitation processes using genetic material from the intervened area,
because native plant individuals species are adapted to the dominant environmental conditions
(Aronson et al., 1993; Cortina et al., 2006; Hobbs and Cramer, 2008).

On the other hand, success of the rehabilitation project depends largely on selected species (Cortina
et al., 2006). Recently, the functional traits framework arises as a useful tool to identify the most
suitable species to rehabilitation goals (Marinez-Garza et al., 2013; Laughlin, 2014; Ostertag et al.,
2015; Engst et al., 2016; Zirbel et al., 2017). Functional traits approach helps us to understand and
predict the responses of species traits to certain environmental conditions and how these traits may
affect ecosystem functions (Lavorel and Garnier, 2002; Zirbel et al., 2017). For example, wood
density has shown a significant and positive relation with resistance to cavitation and specific leaf
area (SLA) has been related to competitive ability and stress tolerance. Species with high SLA have
faster growth rates and are less stress tolerant. Plant height is a predictor of depth root system and
their ability to explore the soil for water. Tall plants present depth roots and dwarf shrubs present
extended and shallow root systems (Weiher et al., 1999; Westoby and Wright, 2006).

The northeast of Patagonia, Argentina, is an ecotonal environment between the Monte and Espinal
phytogeographic regions. From the structural point of view, is a shrubby-steppe with isolated trees
surrounded by tall grasses and herbaceus species. The plant communities are developed in a wide
variety of soils and reliefs (Leon et al., 1998; Bran et al., 2000; Godagnone and Bran, 2009; Morello
et al., 2012; Oyarzabal et al., 2018).

Northeast patagonian ecosystems present obvious signs of degradation and fragmentation (Abraham
et al., 2016; Zeberio, 2012, 2018). The extensive livestock production and rainfed agriculture are
dominant soil uses in this area. Extensive crops are abandoned because of their profitability loss due
to periodic drought (Zeberio et al., 2018). Ecological restoration studies in this area have different
scope and they are not always available (Bran et al., 2007; Funk et al., 2012; Gaitán et al., 2007;
2008; Kropfl et al., 2011; Peter et al., 2012; Zeberio, 2018). Research about the performance of
native woody species in rehabilitation projects in this area are particularly scarce (Torres Robles et
al., 2015; Zeberio et al., 2018). Moreover, we do not know any study that relates functional traits of
these species with their performance in the harsh environmental conditions of degraded northeast
patagonian systems.

We developed Structural Equation Model (SEM) based on hypothesized relationships between

environmental conditions, plant functional traits and species responses. SEM is a useful tool for
understanding the direct and indirect effects of predictors in complex multivariate systems (Grace et
al., 2010).

The aim of this work was to assess the survival rates and growth of native woody species in
degraded areas of the northeast of Patagonia and linked these plant responses to plant functional
traits and environmental variables.

2 Materials and methods

2.1 Site selection

We identify degraded areas in the northeast of Patagonia region based on geomorphological units
defined for Godagnone and Bran (2009). According to Morello et al. (2012), the study area is
located in the eco-region of Monte de Llanuras y Mesetas. This region extends from 32º south
(Mendoza porvince) to 42º (northeast of Chubut province) (Figure 1).

We select sites to develop rehabilitation trials based on two criteria: (1) they have common land use
history (Table 1), but (2) they have differential geomorphological characteristics that contemplate
the high heterogeneity of the area (Table 2). According to these criteria, we select three sites. The
sites were denominated with owner's name. Erripa (Err) and Iturburu (Itu) belong to Adolfo Alsina
district (Río Negro province) and Bosero (Bos) to Patagones district (Buenos Aires province).
Degraded lands represent 40 % of the total surface of these districts (Pezzola and Winschel, 2004;
Zeberio, 2012).

Three plots of 100 m2 were established in each site. The plots were separated from each other by at
least 75 m, and were fenced with a 60 cm high plastic mesh to protect actively planted species
against potential damage caused by livestock and small mammals herbivores.

The climate data were obtained from WorldClim database (Fick and Hijmans, 2017). The mean
annual precipitation of 2013-2016 period presented a bimodal behavior, with two maxims. In the
three sites rainfall reached a peak in April, but reached a second peak in September at Bos site and in
October in the other two sites. Accumulated precipitations at Bos, Itu and Err sites were 1329, 1381
and 1366 mm.yr -1, respectively. In autumn and winter higher rainfalls were registred at Bos and Err
sites. In the spring and summer higher rainfalls were observed at Err site, the closest to the sea coast
(Figure 2). The mean maximum temperature for January (the warmest month) ranged between 27.2
and 25.8 °C and the mean minimum temperature for July (the coldest month) ranged between 0.6
and -0.2 °C according to the site.

Soil chemical properties were estimated from a composite soil sample (30 cm depth) collected to the
centre of each plot. We estimated soil organic matter (MO) by Walkley and Black method, total
nitrogen (N) by micro-Kjeldahl method and total phosphorus (P) by Olsen method. Further, we
registered soil depth in each site until lithic contact and extracted a core to determine apparent soil
density (de Inalbon, 2005).

2.2 Species selection

The selected species were the most frequents in non-degraded areas (Zeberio et al., 2018). Woody
native species selected were shrubs Prosopis flexuosa DC. var. depressa F.A. Roig, Condalia
microphylla Cav. and Schinus johnstonii F. A. Barkley, and the tree Geoffroea decorticans (Gillies
ex Hook. & Arn.) Burkart (Table 3).

Seeds of selected species were collected in the study area in order to produce individuals in a
greenhouse. Seeds were collected from plants that met the selection criteria established for Folliot
and Thames (1983) and were stored in paper bags to sowing time. The sowing of seeds was carried
out between May and July 2012 in the greenhouse of EEA INTA Valle Inferior.

Pre-germination treatments more adequate to each species were applied. Condalia microphylla, G.
decorticans and P. flexuosa seeds were scarified with sulphuric acid (98%P/V) for 32 minutes and in
the case of S. johnstonii only the mesophyll was removed (Zeberio and Calabrese, 2013; Zeberio
and Perez, in press). Seeds were sowing in plastic trays of 60 x 48 x 15 cm contained a mixed
substrate consisting of equal parts of sand and turba, to facilitate removal seedling. After that,
seedlings were disposed in individual pots.

Individual pot contained a substrate consisting of soil extracted from assay sites (50%), commercial
earthworm (25%) and turba (25%). Plants were grown in the greenhouse around seven months with
periodic irrigation (twice a week). After that, rustication process was developed (Cortina et al.,
2006). A month before plants were transplanted to the field, they were gradually exposed to
acclimatation to environmental conditions. Acclimatation implied direct sunlight exposition, wind
exposition and irrigation decrease (once a week).

2.3 Experimental design

Plants were transplanted to the assay sites between April and May 2013. Eleven individuals were
planted in lines, two lines for each species, except for C. microphylla (five plants in one line). This
species presented high mortality in rustication phase and we did not have enough individuals. The
squad of individuals with plants over two years of age was completed. The lines were arranged 2 m
apart from each other. The plant holes were 15 cm in diameter and 20 cm depth, and 1200 cm3 of
commercial earthworm were adding at the bottom of the holes to prevent compaction. The remaining
spaces of the holes were filled with soil extracted when digging the plant hole and were manually
compacted. To improve moisture retention, a basin surround each plant was carried out to capture
and retain natural moisture.

In December 2013 we registered survival of individuals of each species (early survival) and its
height and basal diameter. Three years later, in February 2016, we registered late survival and
measured height and basal diameter of each plant alive. The growth of each plant was estimated as
the difference between the initial and final measurement of the height and basal diameter.

Plant functional traits (specific leaf area, wood density and typical plant height) were obtained from
Try plant trait database (Kattge et al., 2011) (Table 3).

2.4 Data Analysis

Generalized linear models were performed to evaluate plant survival and growth. Binomial
distribution of survival data and normal distribution of growth data was assumed. Site and species
were considered as predictor variable. The interaction site-species was estimated to evaluate
different behaviour of species in each site. We make the comparisons between means using the LSD
Fisher test. Data were analyzed with Infostat software (Di Rienzo et al., 2008).

Causal relationship of environmental variables and plant functional traits to determine their effects
over early survival rates, late survival rates, plant height growth and basal diameter growth were
analyzed. Environmental variables and plant functional traits were selected by linear correlation with
plant survival rates and growth. Structural equation models (SEM) are a synthesis for path analysis,
factorial analysis and maximum likelihood techniques. They are used in ecological science as a
causal inference. This analysis can prove plausibility of causal models based on a priori information
on relationship interest variable. We used SEM to evaluate the relative importance and
direct/indirect effects of climate and soil variables and plant functional traits on survival and growth
(early survival rates, late survival rates, height growth and basal diameter growth). We used the
normed fit index and the root mean square error of approximation index as measures of model fit to
our data (Grace, 2006). Path coefficient estimates were obtained using the maximum-likelihood
estimation technique; they are the equivalent of standardized partial regression coefficients and are
interpreted as relative effects of one variable upon another (Grace 2010).
3 Results

3.1 Early survival

Early survival rate was estimated six months after plants were planted in the restoration trial sites.
Prosopis flexuosa, G. decorticans and S. johnstonii presented high survival rates (near 100%).
Condalia microphylla presented lower survival rates (50%) than the other species (p <0.001) (Figure
3). Bos and Err sites showed higher survival rates (85 and 95%, respectively) than Itu (75%) (p=
0.046). Species*sites interaction of early survival rate was significant only for C. microphylla (p =
0.009). This species presented elevated early survival rates in Err (70%) but lower in Bos (50%) and
Itu sites (20%).

3.2 Late survival

Three years later to transplant date, in summer 2016, late survival rates were registered. Late
survival rates decreased respect to early survival ones. Prosopis flexuosa and S. johnstonii presented
higher late survival rates (70 and 75 %, respectively) than C. microphylla and G. decorticans (15 and
35 %, respectively) (p = 0.0003) (Figure 4).

Late survival at Itu site (60 %) was higher than Bos and Err sites (42 and 38 %, respectively) (p =
0.0074). Species*site interaction was not significant (p = 0.186).

The transplanted specimens of G. decorticans were severely browsed by the Patagonian hare
(Dolichotis patagonum), their stems were felled at ground level.

3.3 Species growth

Three years after transplant, plant height ranged between 25.5 cm (C. microphylla) and 38.54 cm (S.
johnstonii).

Height increase of P. flexuosa (9.76 cm) and S. johnstonii (9.58 cm) was higher than that of G.
decorticans (5.99 cm). Condalia microphylla was the species with the smallest height increase (1.04
cm; p = 0.0002) (Figura 5). Plant height increase was higher at Itu site than Bos an Err sites (p
<0.0001). Species*site interaction was significant (p= 0.047) because G. decorticans and S.
johnstonii presented higher height increase at Itu site than at the other sites.

The basal diameter of stems varied between 0.47 cm (G. decorticans) and 0.96 (S. johntonii). The
plant basal diameter increase of Schinus johnstonii (0.29 cm) and P. flexuosa (0.21 cm) were higher
than that of the other species (p <0.001) (Figura 6). The Itu site presented higher basal diameter
increase than the other two sites (p <0.001). Species*site interaction showed significant differences
(p<0.0001). Schinus johnstonii showed higher increase of basal diameter at Itu site, but C.
microphylla, G. decorticans and P. flexuosa did not presented differences between sites.

3.4 Effects of environmental variables and plant functional traits on survival and growth.

A structural equation model was built to evaluate relations between environmental variables and
functional plant traits over earlier survival rates, late survival rates and growth (Chi square = 43.5; p
= 0,18; D.F = 28; Normed fit index = 0.87). This model explains the 91% of variation in early
survival rates, height growth and diameter growth, and 95% of late survival rates (Figure 7). In the
graphs, only positive and negative statistically significant effects (p <0.05) were presented. The
wood density had a negative relation with late survival rates. The specific leaf area (SLA) showed
positive relation over early survival and late survival rates of transplanted plants to the field. The
typical plant height of the species used in this rehabilitation trial had a negative effect on late
survival rates (Figura 7).

The minimum and maximum temperatures had a negative effect on early survival rates, but they had
a positive effect on late survival rates. Accumulated seasonal precipitation in the study period
positively affected early survival rates, but had a negative effect on late survival rates and height
growth (Figure 7).

Edaphic variables employed in built model which showed statistical significance were organic
matter percent (M.O.) and soil depth to litic contact. Organic matter had a positive effect on late
survival rates, while soil depth had negative effect on early survival rates and height growth (Figure
7).

4 Discussion

The SLA values of the woody species used in this study had a positive impact in their early survival
and late survival rates. It is recognized the existence of two contrasting plants strategies, the
acquisitive, which show a set of functional attributes that allow them a rapid acquisition of resources
and high growth rates. And, on the other hand, the conservative ones, which present another set of
attributes that permit them to keep the acquired resources, but they show lower growth rates
(Poorter, 1990; Lambers and Poorter, 1992; Reich et al., 1997; Aerts and Chapin, 2000; Wright and
Westoby, 2002; Díaz et al., 2004). Plant species with low SLA values tend to have sclerophyllous
leaves. Their thick epidermal walls keep up moisture more efficiently, have lower canopy height,
small xylem vessels that prevent cavitation and high specific wood density (Wright and Westoby,
2002; Díaz et al., 2004; Westoby and Wrigth, 2006). The prevailing plant strategies are in relation to
the environmental characteristics of the region in which they are established (Díaz et al., 2004). In
arid and semi-arid regions of Argentina, the species have relatively low SLA values (Díaz et al.,
1999). This is the case of the species of our study, their leaves have anatomical structures that
protect the photosynthetic tissues from desiccation and herbivory, such as thick cuticles, presence of
spines and anti-herbivore compounds (Bucci et al., 2004; Kraus et al., 2003; Peláez et al., 1994;
Villagra et al., 2011). Therefore, these species may be considered as conservatives ones according to
their SLA low values. These plant conservative strategies could help to achieve high or moderate
survival rates of the selected species in the harsh environmental conditions of the study sites.

Specific leaf area values correlated negatively with wood density values. The wood density is
considered as a proxy of vulnerability to embolism, which can endanger the growth and survival of
plants in water stress conditions. Higher wood density is related to a greater resistance to cavitation
(Westoby and Wrigth, 2006; Chave et al., 2009). Thus, in our study, we expected a positive
relationship between wood density values and survival rates. However, wood density presented a
negative relationship with early survival rates. We believe that this result was conditioned by the
high mortality rate presented by C. microphylla, which have the highest wood density between the
species used in this study. The high mortality of C. microphylla would be related to their
ecophysiological characteristics. Pelaez et al. (1996) reported low C. microphylla survival rates
(25%) in the semi-arid central region of Argentina, where the rainfall shortage impaired the
establishment of transplanted individuals. In our work, it is likely that in addition to environmental
constraints, the older age of transplanted individuals also harmed their survival rate. The specimens
of C. microphylla used in this rehabilitation trial presented high mortality rate in the rustication
phase (unpublished data) and older specimens had to be used to complete the necessary staff. Cortina
et al. (2006) indicate that in order to obtain high survival rates, the specimens of woody species
destined to ecological restoration should not be long-lived and it is desirable that at the time of
transplantation they are about to come in their first vegetative stage.
The typical height of plant species of this study negatively affected late survival rates. This is linked
to the lowest survival rates of G. decorticans, the only tree of this study which is higher than than
that of shrubs.

The plant height growth was related to soil depth. Schenk and Jackson (2005) indicate that woody
plant height is a good predictor of the root system depth. When root systems are limited by some
physical soil impediment, plants have a lower development of aboveground organs too (Bucci et al.,
2004; Peláez et al., 1994). The Bos site had the lowest soil depth due to a sub-superficial calcium
carbonate layer, which acts as a barrier to root system development. The differences in plant height
growth registered between the study sites indicated that soil depth had a sharp effect on transplanted
plants, independently of their species identity. The height growth was higher in plants that were
growing in deeper soil sites.

In our study, P. flexuosa and S. johnstonii were the most successful species, because they evidencied
the highest survival and growth rates. Both species present a dimorphic root system. They have deep
roots that allow them water uptake of deep soil layers, but shallow roots too to take the soil surface
water supported by current rainfalls (Bucci et al., 2009; Jobbágy et al., 2011; Villagra, 2000;
Villagra et al., 2011). High early survival rate of P. flexuosa obtained in this study was similar to that
obtained by López Launstein et al. (2012) in the northwest of Argentina. In that study, P. flexuosa
showed 100% of early survival rates under moderate water stress, and 70% of survival rates under
severe water stress, in field conditions. Pérez (2013) in the northwest of Patagonia region registered
survival rate in S. johnstonii (75%) lower that obtained in our study. Both species, P. flexuosa and S.
johnstonii, were different in SLA values, but similar in wood density and root system morphology.
Dimorphic root systems are absent in the other species of this rehabilitation trial. Condalia
microphylla presents a shallow radical system, extended in the first 60 cm soil depth, which makes it
to compete with grasses to obtain rainfall water (Peláez et al., 1994). Instead, G. decorticans present
a pivoting and deep root system (Krauss et al., 2013), which makes it less able to take advantage of
superficial soil water. The dimorphic root system of P. flexuosa and S. johnstonii, allow them to
obtain water of superficial and deep soil layers. This may help to understand the persistence of these
species under water stress conditions (Díaz and Cabido, 1997; Whitford, 2002). On the other hand,
P. flexuosa and S. johnstonii are shrubs, while G. decorticans are tree, but plants of the last species
were severely browsed by Patagonian hare in the field, which does not allow us to obtain conclusive
data. For this reason, the growth form (trees vs. shrubs) would not be sufficient to explain the
success of shrubs in our study.

The maximum and minimum temperatures may be indicators of the effects of extreme temperature
on plants. These temperatures presented a negative effect on early survival rates of plant species in
this study. Instead, temperatures showed a positive effect on late survival rates. These results would
be associated with the critical threshold that plants must be overcome for the first year after
transplanted to the field and rustication process (Maestre et al., 2006; Cortina et al., 2006).

Rainfall had a positive effect on early survival rates, but a negative effect on late survival rates and
plant height growth in all transplanted individuals of our study. Northeast of patagonian region is
characterized by the inter-annual and intra-annual rainfall variability (Godagnone and Bran, 2009).
For this study measurement period, the rainiest seasons were autumn (80 mm) and spring (71 mm),
and the lowest one was the summer (25 mm). Early survival of plants was estimated in spring, one of
the rainiest period, but three years later, plants have been exposed to summer rainfall shortages. The
plant transpiration demands are the highest during the summer and the drought could have
negatively affected late survival rates and limited plant growth.
Soil organic matter of each of the studied sites were in the range of that reported by Godagnone and
Bran (2009) (0.46% to 1.63%) for northeast of Patagonia region. However, this low soil organic
matter values had a positive effect on late survival rates of plant species in our study. This might be
related to native species adaptation to obtain scarce soil resources and use them efficiently. Also, the
sites of this study presented a common land use history, which was rainfed agriculture. Grman et al.
(2013) found that residual nutrients from previous crop land use may have a positive effect on
survival and establishment of plant in restoration areas.

5 Conclusions

The structural equation models used in this study related the performance of native woody species
with functional traits and environmental variables in an ecological restoration trial in deforested
semiarid lands.

Prosopis flexuosa and S. johnstonii were the most successful species of our study. These species
overcome rainfall shortage and high temperatures periods. It is possible that the presence of a
dimorphic root system allows them to have competitive advantages regardless of the observed
functional traits. These advantages would be related to the use of deep soil water as the soil water of
shallower layers.

Due to the damage caused by the browsing of the Patagonia hare, it is desirable to continue with
field studies of the survival and growth of G. decorticans to evaluate their inclusion in ecological
rehabilitation projects in deforested areas of the northeast of Patagonia.

Condalia microphylla is a frequent species in northern Patagonian region and is desirable intensify
their autecology studies to include this species in rehabilitation project.

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Table 1. Rehabilitation trial sites location and land use in each one.

Sitio Latitud Longitud Use Age of clearing

Er 41º 9' 11,86''S 63º 25' 19''O Rainfed agriculture 25
It 41º 1' 4''S 63º 1' 40,3''O Extensive livestook 23
Bo 40º 39' 30,4'' S 62º 52' 0,1'' O Rainfed agriculture 20

Table 2. Characteristics of Rehabilitation trial sites EC: Electrical conductivity. BD: Bulk density. O.M.: Organic
Matter. N: Total nitrogen in percentage. P (ppm): Phosporus in ppm.

Site Geomorphological region Soil texture Depth pH EC BD MO N P

Err Planicie loessica Sandy >1 8.2 0.58 1.27 0.53 0.06 8.93
Itu Planicie loessica Sandy- loamy 0,45 8.2 0.54 1.41 1.,06 0.09 8.77
Bos Planicie interfluviales mesetiformes Loamy 0.12 7.8 1.78 0.43 0.300 0.03 2.31

Table 3. Summary of plant characteristics used in the rehabilitation trials. SLA: specific leaf area (cm2.gr-1). WD:
Wood density (gr.cm2)-1. P H: Typical Plant Heigth (m).

SLA WD Foliage PH Life form

C. microphylla 26.5 1270 Evergreen 0.5- 2 Shrub

G. decorticans 103.2 970 Deciduous 1.4- 4 Tree

S. johnstonii 28.5 995 Evergreen 1.5 Shrub

P. flexuosa 141.5 980 Deciduous 2.5 Shrub

Figure 1. Study area and rehabilitation trial sites location.
Figure 2. Rainfalls occurred between 2013-2016 in Bos, Itu and Err sites and historical monthly average precipitation
(data extracted to de Berazategui, 2004).
 A A
A A
A A A
100,00
Bos
A
A Itu
90,00
Err
A
80,00 B

70,00
Early survival (%)

60,00

50,00
B
40,00

30,00

20,00

10,00

0,00
C. microphylla P. flexuosa S. johnstonii G. decorticans
Species

Figure 3. Woody species early survival rates recorded in January 2013 (seven months from field planting). Means
with a different letter indicate significant differences between sites for each species (p <0.05).

A
100
ABC Bos
90 AB BCDE Itu
Err
80
BCD
70
BCDE
Late survival (%)

60
CDE

50

DE
40 DE
CDE
30
DE
20 E

10

0
C. microphylla P. flexuosa S. johnstonii G. decorticans
Species

Figure 4. Woody species late survival rates recorded in January 2016 (31 months from field planting). Means with a
different letter indicate significant differences between sites for each species (p <0.05).
 40,00 Bos

A DEF
Itu
35,00 Err

30,00
B BCDE
BC EF
25,00
Height (cm)

BCD
BCDEF
20,00
CDEF

15,00

10,00
DEF
EF F
5,00

0,00
C. microphylla P. flexuosa S. johnstonii G. decorticans
Species

Figura 5. Woody species height growth (cm) between 2013-2016. Means with a different letter indicate significant
differences between sites for each species (p <0.05).

1,50 Bos
1,40 Itu
1,30 Err
1,20 A
1,10
1,00
0,90
Diameter (cm)

0,80
0,70
0,60 B
B BC
0,50 BCD

0,40 BCD BCD BCD

0,30
D CD
0,20 CD
0,10 CD

0,00
C. microphylla P. flexuosa S. johnstonii G. decorticans
Species

Figura 6. Woody species basal diameter growth (cm) between 2013-2016. Means with a different letter indicate
significant differences between sites for each species (p <0.05).
Figure 7. Structural equation models for the environmental variables (mean minimum temperature of July, T°min
July; º mean maximum temperature of January, T° max January; accumulated precipitation of the season,
Accumulated Pp; Soil Depht and soil organic matter percentage (MO %), plants functional traits (specific leaf area,
SLA; wood density; typical plant height) and estimated variables (early survival rates; late survival rates; diameter
growth; height growth). The numbers adjacent to arrows are path coefficients and they show the strengths of the
effect. Non-significant (p > 0·05) paths were eliminated. Significance levels are as follows: **p < 0·01; ***p <
0·001.