Journal of Arid Environments 161 (2019) 1–10

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Journal of Arid Environments

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Towards a dryland framework species approach. Research in progress in the T

Monte Austral of Argentina
Daniel R. Péreza,∗, Fernando M. Farinaccioa, James Aronsonb
a
Laboratory of Rehabilitation and Ecological Restoration of Arid and Semiarid Ecosystems (LARREA), National University of Comahue, Buenos Aires 1400, CP 8300,
Neuquén, Argentina
b
Center for Conservation and Sustainable Development, Missouri Botanical Garden, 4344 Shaw Blvd, St Louis, MO, 63110, USA

A R T I C LE I N FO A B S T R A C T

Keywords: We discuss the strategies and early results from an ecological restoration and rehabilitation project underway in
Arid and semiarid Monte Austral (Southern Monte) of Argentinian Patagonia. We use this as a platform to propose a dryland
Attributes framework species approach (DFSA) to ecological restoration and rehabilitation (ERR). DFSA is based on the
Desertification ‘framework species approach’ developed in mega-diverse tropical forest biomes of Australia and southern Asia
Species selection
over the past 20 years. We discuss how the approach could be adapted and modified to boost the impact of ERR
and help achieve more effective recovery of biodiversity, ecosystem health, and human health and well-being in
the vast dryland ecosystems of the world.
In arid and semiarid ecosystems, not all native species respond well to methods for growing in nurserys, and a
key to ERR success is the identification of a coherent set of species selected on the basis of multiple criteria, both
ecological and practical, and simple tests for field performance. We describe selection criteria and early results of
field trials of three candidate framework plant species undergoing field trials in our region. Of these three,
Atriplex lampa and Hyalis argentea var. latisquama showed uniformly high performance while the response of
Senecio subulatus var. subulatus varied according to substrates. We also discuss the use of drones, and other tools
for both research and action in conjunction with DFSA.

1. Introduction urban and rural.

Ecological restoration and rehabilitation (hereafter, ERR) are key
strategies to cope with, and at least partially reverse, the effects of
desertification and ecosystem degradation in the vast arid and semiarid
areas (hereafter Drylands), home to approximately 2.5 billion people,
mostly in developing countries (UNCCD, 2011). Worldwide, degrada-
tion and desertification is detrimental to ecosystem health and human
health in drylands (James et al., 2013). Moreover, the re-establishment
of vegetation in severely degraded drylands is slow and, in regions like
our study area in Argentina, unassisted recovery can take centuries.
What's more, it will probably never happen without intentional re-
storative interventions, such as substrate amelioration and subsequent
sowing of seeds and/or planting of seedlings (Bainbridge, 2007; Abella
et al., 2015). Finally, drylands are every bit as hard hit by anthro-
pogenic climate disruption as coastal areas or forested biomes are
(World Bank and UNCCD, 2016). Hence, the stakes are high vis à vis
biodiversity, mitigating and adapting to climate perturbations, and
optimizing the flow of ecosystem goods and services to all people,
Regardless of the underlying drivers of desertification in a given
area, effective interventions for dryland restoration and rehabilitation
must focus both on biotic and abiotic obstacles, and use native plants
found to be helpful to stop degradation and promote the recovery of the
structure, composition, and functionality of impaired ecosystems
(Milton et al., 1994; Whisenant, 1999). Nevertheless, despite decades of
research (e.g., Noy-Meir, 1973), advances in understanding the ecology
of dryland biomes have not yielded proportional advances in our ability
to restore degraded dryland ecosystems (James et al., 2013). Likewise,
the development of methods for assessing the success of the actions to
combat desertification is recognized as a top priority among the sci-
entific community (Costantini et al., 2016). In this context, we propose
a new approach to restore and rehabilitate dryland ecosystems, and we
present initial results that show that its application is possible. By
borrowing and adapting some aspects of the original Framework Spe-
cies Approach (Goosem and Tucker, 1995) developed for application in
mega-diverse, humid tropical forests (Goosem and Tucker, 2013), it
may be possible to expand and improve the effectiveness and longevity

∗
Corresponding author.
E-mail addresses: danielrneuquen@gmail.com (D.R. Pérez), fer.farinaccio@gmail.com (F.M. Farinaccio), james.aronson@mobot.org (J. Aronson).

https://doi.org/10.1016/j.jaridenv.2018.09.001
Received 15 February 2018; Received in revised form 1 September 2018; Accepted 3 September 2018
Available online 11 September 2018
0140-1963/ © 2018 Elsevier Ltd. All rights reserved.
D.R. Pérez et al.
Fig. 1. Study area. A: location of the Monte, the Monte Austral (Southern Monte), and planting sites (S1, S2, S3, S4, S5). B: Neuquén Province, Monte and Monte
Austral in the Argentinean Republic (Geographical coordinates system WGS-84).
of restorative activities in drylands.
The initial idea of the framework species method or approach was to
establish particular mixtures of species which act as ecosystem building
blocks and attract seed dispersing wildlife. Wild animals, attracted by
the planted trees, disperse the seeds of additional species into planted
areas, while the cooler, more humid, and weed-free conditions, created
by the planted trees, favour seed germination and seedling establish-
ment (Goosem and Tucker 1995, 2013; Elliott et al., 2013).
While in humid tropical forests, such as those where the pioneers of
the Framework Species Approach have worked, the practical con-
straints to overcome have to do in part with the huge number of native
plant species, and the need to identify the most suitable ones for use in
jumpstarting ecological succession in open sites, several different pro-
blems must be faced is drylands. For example, far more than in wet
tropical regions, the survival and development of plants in degraded
drylands depend in large measure on the specific soils in which they are
planted or sown (Costantini et al., 2016). At the same time, numerous
studies show that the vigor and resistance of seedlings and their ability
2
Journal of Arid Environments 161 (2019) 1–10
to establish and survive in severely degraded dryland sites is also re-
lated to the functional groups (e.g., grasses, forbs, shrubs, etc.) to which
they belong, as well as the quality of nursery plants and plantation
techniques, and of course herbivory, among other factors (Bainbridge,
2007; Abella and Newton, 2009; Commander et al., 2013; Yirdaw et al.,
2017; González and Pérez, 2017). These factors are not unique to
drylands but they do add up to a kind of early ‘bottleneck’ for effective
restoration and rehabilitation at large scales.
Therefore, restorationists in drylands need to identify which among
native plant species, and functional groups of species, are most likely to
show the best growth and adaptability under a restoration context. This
is not trivial: the spectra of plant life forms in arid lands are among the
most diverse of all terrestrial biomes on Earth (Smith et al., 1997), and
to achieve restoration ideally they should all be represented. Faced with
these problems, there are monumental challenges and great opportu-
nities related to the process of developing an effective Framework
Species Approach (FSA) applicable to drylands, which we will call
Dryland Framework Species Approach, or DFSA.
D.R. Pérez et al.
The modifications and adaptation of the original framework species
approach that we present here are not without precedent. For example,
FSA was effectively adapted to restoration of seasonally dry tropical
forest (Elliott et al., 2003), over-exploited agricultural sites
(Wangpakapattanawong and Elliot, 2008), and bauxite mine tailings
(Corrêa Dias et al., 2014). Additional examples of particular applica-
tions and elaborations to specific contexts include the preferential use
of fire-resistant species (Elliott et al., 2003), linking FSA with efforts
aimed at assisting spontaneous regeneration, and careful consideration
of local traditional ecological knowledge and customs (Elliott et al.,
2013).
The seven attributes of plant species used by Goosem and Tucker
(2013) in their selection process for the FSA were, in order of im-
portance (and in our own words): a) Tolerance of open conditions; b)
Attractiveness to fauna, especially seed-dispersing birds and mammals;
c) Early production of wildlife resources; d) Presumed role as ‘Keystone’
species; e) Ability to help create new habitat away from planting sites;
f) Rapid or persistent growth; and g) Easy and relatively rapid germi-
nation.
In the DFSA that we propose here, we emphasize four concerns to
consider, as follows: 1) Species selection; 2) Local knowledge; 3)
Differing spatial layouts and mechanically engineered micro sites, both
near and far from planting sites; 4) Efficient monitoring from drones for
adaptive management); and three attributes: a) High survival and
growth rates; b) Attractiveness for fauna; and c) Easy to germinate.
In this paper we present current results on relevant ecological and
silvicultural traits of three candidate native plant species for DFSA se-
lected for initial trials among 22 species under consideration in
Patagonia. We address the first two concerns and discuss future appli-
cations of the other two as well as the three attributes for DFSA ap-
plications.
2. Materials and methods
2.1. Study area
The present study was conducted in the most arid region of
Argentina, the so-called Monte, specifically in the “Monte Austral
Neuquino” (Busso and Fernández, 2017; Busso and Bonvissuto, 2009),
in Neuquén Province, hereafter “Monte Austral” (Fig. 1). Vegetation is
generally distributed in oval patches of approximately 300 cm x
200 cm, formed by shrubs and grasses with high horizontal and vertical
heterogeneity (Busso and Bonvissuto, 2009). The most abundant shrub
species are Larrea divaricata Cav., L. cuneifolia Cav., L. nitida Cav.,
Monttea aphylla (Miers.) Benth. & Hook. var. aphylla, Bougainvillea spi-
nosa (Cav.) Heimerl, and Atriplex lampa (Moq.) D. Dietr. Common
herbaceous plants include the perennial grasses Pappostipa speciosa var.
speciosa (Trin. & Rupr.) Romasch, Leymus erianthus (Phil.) Dubcovsky,
and Poa ligularis Nees Ap. Steudel, all typically found growing under
shrub canopies. In addition, vegetation on clayey and saline soils is
generally dominated by salt-tolerant Chenopodiaceae and Halophytaceae
in the genera Atriplex, Suaeda and Halophytum (Busso and Bonvissuto,
2009).
With respect to fauna, there is a large guild of granivorous birds,
large herbivores and rodents that interact with the vegetation in the
Monte (Milesi et al., 2002; Campos and Velez, 2015). Ants are common,
and several species can transport fruit segments, scattering seeds on
trails an around ant nests (Pirk et al., 2009).
The Monte Austral has mean annual temperature of 12 °C; the
lowest temperature ever recorded was −17 °C. The ratio between
precipitation and potential evapotranspiration ranges between 0.05 and
0.5, indicating a marked water deficit, and the annual rainfall is highly
variable, and occurs mainly in winter or early spring May to October
(Labraga and Villalba, 2009). Between 2003 and 2013, rainfall varied
between 40.4 and 288.8 mm per year in our study area (AIC, 2017).
November, December, and January (mid-spring to early summer) have
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Journal of Arid Environments 161 (2019) 1–10
the greatest wind speeds, with a mean of 17 km per hour at 2 m above
ground level (Busso and Bonvissuto, 2009). Prevailing soils are aridisols
and entisols, with an aridic hydrothermal regime, high pH, and CO3,
and low levels of organic matter (Busso and Fernández, 2017). The
main anthropogenic disturbance factors are domestic animal grazing,
pastoral fires, gas and oil exploration and extraction, and extraction of
fuelwood, all of which began to have serious impact starting two cen-
turies ago, linked to European colonization (Villagra et al., 2009; Busso
and Fernández, 2017).Of this vast dryland region, which as a whole
comprises approximately 78.5 million ha, no less than 73.5 million ha
(93.6%) are considered mildly to severely desertified (Mazzoni and
Vazquez, 2009). In short, desertification appears to be irreversible in
the absence of active interventions (Abella, 2010; González and Pérez,
2017).
2.2. Soil and climate in experimental sites
Plantations of native species were carried out in five sites 0.7–1 ha.
in size (S1 – S5, Fig. 1), separated by distances between 1 and 5 km.
Altitude varied between 394 and 469 m.a.s.l. The sites were all aban-
doned oil platforms that lacked vegetation cover. Although the sites had
a history of oil exploration, there were no residues of hydrocarbons in
the soil profiles. Rainfall was recorded once a month in a direct-reading
polypropylene pluviometer placed in each plantation site. Soil physical
and chemical properties in each site were assessed through samples
collected to 50-cm depth. Each sample was composed of ten random
soil sub-samples. Soil temperatures were registred with data loggers
(Sensors Thermochron iButton®). Soil surface moisture in three random
sampling locations was measured between 40 and 60-cm depth through
direct reflectrometry measurements with a TDR probe (Time Domain
Reflectometry-Fieldscout model 300). Water infiltration was assessed in
a tension water infiltrometer (Decagon Devices-model S). and soil
compaction to a depth of 20 cm was measured with the aid of a punch
manual penetrometer (2,5 kg/cm2 of load). Ph was assessed through the
RLSAV procedure; soil electrical conductivity and sodium load were
measured through conductometry applied to saturated soil paste and
sodium absorption ratio (SAR), respectively.
2.3. Concern 1: species selection
Plant species growing and reproducing under very stressful en-
vironmental conditions were the “potential” or primary candidates for
inclusion in our experimental framework species group. These extreme
conditions occur in various highly degraded states caused by livestock
overstocking or vegetation clearing, exacerbated by periodic rain-
storms, combined with sheet runoff over impermeable rocky substrates
and steep slopes. The chosen sampling sites in this work were margins
of temporary streams (TS), sites with steep slopes of approximately 40°
(S), and open sites of degraded Monte vegetation with less than 30%
plant cover (MM). The line-intercept method was used to record species
richness along three transects of 50 m in each habitat type (Matteucci
and Colma, 1982).
2.4. Concern 2: local knowledge
In this first application of DFSA, with regards local knowledge, we
considered only the indispensable need for local people to have the
capacity to produce at least 5000 viable seedlings of each of the puta-
tive framework species. In arid zones such as those of Patagonia, there
is a dire lack of knowledge about the production of seedlings for re-
storation due to the historical devaluation of this type of environment
in which only extensive cattle ranching and oil exploitation take place
(Pérez et al., 2017). Other related aspects that need to be considered
such as local input into species selection are mentioned in the discus-
sion and conclusions sections below.
D.R. Pérez et al. Journal of Arid Environments 161 (2019) 1–10

2.5. Attribute a: survival and growth 2.7. Attribute c: easy to germinate

A total of 2,500 seedlings for each of the three selected species were
planted in April–May 2012, at age 9 months. They were randomly se-
lected from a pool of 15,000 individuals (5,000 seedlings of each spe-
cies) produced in the nursery garden of the National University of
Comahue in plastic pots (270 cm3) filled with native soil, perlite, and
organic compost mixture. Five hundred seedlings from each of the three
species were planted in each of the five different sites. The area where
the sites are located is indicated in Fig. 1. Planting holes were made
with a hand-held soil hole drilling machine (Seery Model HT10®); to
make holes of 40 cm depth and 20 cm diameter, in which 500 ml of
polyacrylamide hydrogel (GELFOREST®) and 1 liter of water were to be
added in each hole. No additional irrigation was carried out. All seed-
lings were protected against herbivorous mammals by a 30-cm high
metallic mesh protection barrier. The five hundred plants per species
were planted in lines 1 m apart, with 1 m spacing between individuals
in each site (S1-S5). A total of 50 specimens were chosen at random for
each species in each treatment in order to assess rates of survival 11 and
22 months after planting (March 2013 and March 2014, respectively).
These dates correspond to the end of summer, which is the most critical
season for plant survival in our study area as ground temperatures can
reach 65 °C (Table 1).
Early plant growth performance was assessed by recording height
and biomass of seedlings before planting, and after 11 months, was
destructive harvest of 15 randomly selected individual was carried out.
Seedling height (cm) and stem diameter (mm) were measured with a
millimeter ruler and an electronic caliper, respectively. Shoots (stem
and leaves) were separated from roots, roots were rinsed thoroughly
running water, and roots and shoots were oven-dried for 24 h at 70 °C.
Next, every section was weighed on analytical scale (0.001 gr).
Likewise, 22 months after planting, the crown area of 50 randomly
chosen seedlings of each species in each site was measured by oval area
(r1 –major radius (cm) - *r2 –minor radius (cm)- *π), and then com-
pared with the crown areas of individuals belonging to the same, lo-
cated 100–500 m from each plantation site (S1 to S5).
2.6. Attribute b: attractiveness for fauna
Attractiveness for fauna and easy to germinate was studied by
bibliographic review carried out using Google Scholar and by con-
sulting websites of all the relevant research centers in the arid zones of
Argentina. Due to the fact that there are few works for the Monte
Austral that address the relationship between fauna and native plants,
consultations were also made with local residents.
The germination capacity of the species was evaluated in the sci-
entific literature. We evaluate whether expensive pre-germination
treatments permit higher levels of seedling production, in local nur-
series, as compared to procedures without pre-treatments. It is im-
portant to consider that germination issues are not only important for
nursery production of the framework species. Seeds of native species
that germinate quickly and easily may have a better chance of getting
well-established thanks to the enhanced micro-environmental condi-
tions provided by each well growing tree in the experimental planta-
tions. Thus, survival and growth (attribute a), attractiveness for fauna
(attribute b), and now attribute (c) of our proposed DFSA are inter-
dependent and complementary in their respective contributions to
achieving effective dryland restoration.
2.8. Statistical analysis
Pearson's linear correlation coefficient was applied to assess re-
lationships between survival rate and soil infiltration, and between
survival rate and soil compaction. A generalized linear model for binary
data was used to assess plant survival. The significance level was set at
5% for all tests, and R software version 3.2.2 (R Core Team, 2015) was
used to perform all analyses. Differences in growth and biomass were
analyzed by paired t-tests.
3. Results
3.1. Concern 1: species selection
A total of 24 plant species initially considered to have high potential
in a FSA context were found growing in the Monte (MM), Temporary
streams (TS), and Steep slopes (S) habitats (Table 2). One of these,
Habranthus jamesonii (Baker) Ravenna, was eliminated because it is
rare, and large-scale propagation was impossible and, secondly, Bromus
tectorum L. (cheatgrass) was rejected as a noxious, non-native weed, so
the number of potential species dropped to 22.
3.2. Concern 2: local knowledge
Only three of the 22 native species had the necessary characteristics
of high production in nursery gardens, namely Senecio subulatus D. Don
ex Hook. & Arn. var. subulatus (Asteraceae) Hyalis argentea D. Don var.
latisquama Cabrera (Asteraceae), and Atriplex lampa (Moq.) D. Dietr.
(Chenopodiaceae). The first is a perennial shrub, the second is a her-
baceous perennial, and the third is a shrub. Environmental education
for adults implemented in conjunction with the participatory action
research also allowed the sharing of knowledge among researchers and

Table 1
Summary of environmental variables in the 5 planting sites, S1-S5. The data are presented as mean ± standard deviation (SD).
Environmental parameter Measuring unit Planting sites

S1 S2 S3 S4 S5

Precipitation (mm) 11 months Total accumulation 145 147 148 143 145
Precipitation (mm) 22 months 119 124 117 120 118
Soil temperature (°C) Maximum 56.5 60 60 61 65
Minimum −4.5 −3 −2.5 −1.5 −3.5
Average 20 ( ± 7) 21 ( ± 7) 22 ( ± 7) 22 ( ± 6) 20 ( ± 6)
Soil moisture (%) On surface 21 ( ± 8) 19 ( ± 5) 24 ( ± 7) 26 ( ± 9) 31 ( ± 13)
In deep (40–60 cm) 14 ( ± 7) 11( ± 2) 18 ( ± 8) 20 ( ± 11) 25 ( ± 18)
Infiltration (mm/min) —————————————————————— 66 ( ± 22) 84 ( ± 8) 43 ( ± 27) 63 ( ± 45) 19 ( ± 8)
Soil hardness (N° hits) Up to 40 cm 91 ( ± 64) 82 ( ± 72) 41 ( ± 35) 38 ( ± 32) 88 ( ± 70)
pH of the soil Sample composed of 10 sub-samples 7.6 7.8 7.5 7.5 7.8
C.E. of the soil (dS/m) 4.62 3.02 21.2 13.3 2.5
RAS 1.1 5.7 4.6 26.1 2.8

4
D.R. Pérez et al. Journal of Arid Environments 161 (2019) 1–10

Table 2 Height was significantly different between time of outplanting and

List of species and families found in habitats MM (Monte), TS (Temporary age eleven months in all three species assessed (p < 0.001). A. lampa
streams), and S (Steep slopes of 40° or more). Each species is categorized as increased in size 3.82-fold during this period, while S. subulatus var.
belonging to one of the following life forms: PS: Perennial shrub, AG: Annual subulatus grew 2.64-fold, and H. argentea var. latisquama grew 3.62-fold.
grass, SP: Succulent perennial sub-shrub, PG: Perennial grass, PF: Perennial
The length of the longest horizontal branch of this latter species was
forb.
considered to be equivalent to height in the other species, since it is a
Species/Family Life form Habitat creeping perennial rather than upright like the other two species
(Fig. 4).
MM TS S
Additionally, there were significant differences in growth rates in
Amaryllidaceae the shoot and root biomasses of A. lampa, S. subulatus var. subulatus, and
Habranthus jamesonii (Baker) Ravenna PG x x H. argentea var. latisquama (p < 0.001). Eleven months after out-
Anacardiaceae planting, shoot and root biomass in A. lampa had increased 3.83-fold
Schinus johnstonii F.A. Barkley PS x
and 3.25-fold, respectively, as compared to 3.44- and 4.73-fold in S.
Asteraceae
Chuquiraga erinacea D. Don PS x subulatus var. subulatus, and 2.8- and 3.61-fold in H. argentea var. la-
Chuquiraga rosulata Gaspar PS x x tisquama (Fig. 4).
Gutierrezia solbrigii Cabrera PS x x x There were no significant differences between the crown area of
Hyalis argentea D. Don ex Hook. & Arn. var. latisquama PG x x
seedlings in planted sites and that of individuals growing in the re-
Cabrera
Senecio subulatus D. Don ex Hook. & Arn. var. subulatus PS x
ference ecosystem (p > 0.10). A. lampa showed the highest variability
Cactaceae of mean crown area of individuals sampled in plantations, and those
Echinopsis leucantha (Gillies ex Salm-Dyck) Walp. SP x sampled in the reference ecosystem site (Fig. 4).
Chenopodiaceae
Atriplex lampa (Moq.) D. Dietr. PS x x
3.4. Attribute b: attractiveness for fauna
Fabaceae
Adesmia guttulifera Sandwith PS x
Cercidium praecox (Ruiz & Pav. ex Hook.) Harms ssp. PS x x x Table 3 shows the results of a literature review on interactions of the
glaucum (Cav.) Burkart & Carter three candidate plant species and various groups of native vertebrates,
Hoffmannseggia glauca (Ortega) Eifert PG x x x including seed-eating birds, large and medium herbivores, small ro-
Senna aphylla (Cav.) H.S. Irwin & Barneby PS x x x
Prosopis flexuosa DC. var. depressa F.A. Roig PS x x x
dents, as well as domestic herbivores, mostly goats and cows. Stem and
Malvaceae leaf consumption was the most common use made by all groups of
Lecanophora ecristata (A. Gray) Krapov. PG x vertebrates, followed by fruit consumption. Many animal species also
Nyctaginaceae use the three candidate framework species for nesting and resting.
Boungainvillea spinosa (Cav.) Heimerl PS x x x
Local inhabitants mentioned the influx of large numbers of grani-
Plantaginaceae
Monttea aphylla (Miers) Benth. & Hook. var. aphylla PS x x vorous birds affecting all three planted species. It was also mentioned
Poaceae by rural residents that rodents are commonly observed in communities
Bromus tectorum L. AG x with a high dominance of A. lampa. In one of the plantation sites, in the
Pappostipa speciosa (Trin. & Rupr.) Romasch. PG x x second year post-plantation, a nest of the granivorous bird Eudromia
Solanaceae
Lycium chilense Miers ex Bertero var. chilense PS x x x
elegans (Elegant crested tinamou) was observed by the authors under
Verbenaceae the cover of planted H. argentea. This indicates the fitness of at least one
Acantholippia seriphioides (A. Gray) Moldenke PS x x of the candidate plant species to become a nesting refuge for local
Mulguraea ligustrina (Lag.) N. O'Leary & P. Peralta var. PS x avifauna. Also all local inhabitants agree that all three candidate species
ligustrina
are desired by livestock as fodder albeit in different phenological stages.
Zygophyllaceae
Larrea cuneifolia Cav. PS x x x
Larrea divaricata Cav. PS x x x 3.5. Attribute c: easy to germinate

The seeds of all the three species require little or no pre-germination

local people that made it possible to produce these and other candidate
species for ecological restoration experiments in three localities in the
Monte Austral Neuquino, since 2010. Participants learned how to col-
lect seeds, how to prepare potting soils for seedlings, and the complete
process of plant production and planting with the support of the
National University of Comahue (Pérez et al., 2017) (Fig. 2).
treatment. Hyalis argentea var. latisquama, has high germination values
without pre-treatment (Camina et al., 2013). Atriplex lampa seeds need
be placed for three days in running water to allow germination. Senecio
subulatus var. subulatus germinate without pre-germination treatments.
4. Discussion

3.3. Attribute a: survival and growth (shoot length, above- and below-
ground biomass, crown area)
Survival rate was similar for all three species eleven months after
planting (p = 0.98); mean A. lampa survival was 91% ( ± 8.4), H. ar-
gentea var. latisquama was 89% ( ± 5.0), and S. subulatus var. subulatus
was 84% ( ± 11) (Fig. 3). After 22 months following outplanting, sur-
vival was significantly different among species (p = 0.003). Specifi-
cally, survival rate was the same in A. lampa (91% ( ± 8) and H. ar-
gentea var. latisquama, (89% ± 5.0) but reduced and showing large
standard deviations in S. subulatus var. subulatus (32% ± 27) (Fig. 3).
Precipitation and soil characteristics in the five study sites are
presented in Table 1. S. subulatus var. subulatus showed positive cor-
relation between soil hardness and survival (R2 = 0.98; p < 0.01; S.
subulatus var. subulatus survival = −12.38 + 3.03 x hardness).
4.1. Concerns for DFSA
4.1.1. Concern 1: species selection
The reduction to a limited number of species to use is a character-
istic of this methodological approach. According to Elliot et al. (2013)
in a first stage preliminary screening, based on current knowledge, the
first step is to identify ‘candidate’ framework species and then embark
on nursery and field experiments with the candidate species. Initially,
detailed information about each species is likely to be sparse. As the
results of nursery experiments and field trials gradually accumulate, the
list of acceptable framework species can be gradually refined.
In our study, we carried out a preliminary examination of species
that are found in abundance in habitats with high environmental stress
as candidates for DFSA. It is important to point out that many of these
species that were able to survive in the three environments surveyed

5
D.R. Pérez et al.
Fig. 2. (A) Members of the native drylands species nursery of the ecological restoration cooperative called “Atriplex lampa”, engaged in plant production tasks;
Neuquén, Patagonia Argentina; (B) Before stage of planting in one of the plantation sites; (C) Placement of protection meshes against herbivores during planting; (D)
View at 22 months after plantation. Photo credits: A) Florencia del Mar González; B) and C) Daniel R. Pérez; D) Fernando Farinaccio.
(cited in Table 2) could not be produced by nurseries due to lack of
local knowledge to overcome mortality problems or difficulties in
achieving the germination of large numbers of seeds. The high mor-
tality recorded for S. subulatus var. subulatus in the second year after
outplanting underlines the importance of gathering and processing all
available ecological and ecophysiological information for each candi-
date species. For example, studies of infiltration and nutrient cycling
may be useful aids to decision-making in species selection for restora-
tion efforts (Costantini et al., 2016). Similarly, knowledge of capacity
for drought tolerance and patterns of resource allocation in the various
candidate species planted together can be crucial (Fernandez et al.,
2015; González-Paleo and Ravetta, 2013).
To illustrate similarities and differences in the application of FSA in
different ecosystems, recall that in the Goosem and Tucker's restoration
work in humid tropical forests of Queensland, Australia, between 30
and 40 tree species of varying successional stages were planted
(Goosem and Tucker, 2013). This represented just a small fraction of
the total number of tree species found in intact forests in the area. Even
in a seasonally dry tropical forest, with lower tree species diversity
Elliot et al. (2003) found that among the 37 species studied, only nine
could be ranked as 'excellent' candidates. In drylands, there is typically
an order of magnitude lower woody plant species richness than is found
in tropical forests. Further, based on vast experience in plant nurseries
in many drylands, it is likely that only a small portion of the woody
species found in intact reference ecosystems will be easy to propagate
and reintroduce successfully. Therefore, we hypothesize that in most
dryland areas, typical characterized by low plant diversity, generally
high environmental stress for plants, and large differences in plants'
growth patterns and adaptations to soils and extreme conditions, the
number of species appropriate for use with a DFSA strategy in fact may
be quite few.
6
Journal of Arid Environments 161 (2019) 1–10
4.1.2. Concern 2: local knowledge
In our work, we tested the possibility that Senecio subulatus var.
subulatus, Atriplex lampa, and Hyalis argentea var. latisquama could be
produced in local nurseries, and evaluated their overall respective
performance in the first two years of growth. However, successful ap-
plication of DFSA in Monte Austral and other drylands will also require
empowerment and active participation of local people whose lands and
livelihoods will be affected by the process and outcome of restoration
efforts. The results that arise from the analysis of the most appropriate
species to tolerate the most adverse environmental conditions in our
study area (concern 1) and concerns 3 and 4, below, must be integrated
with the help of local knowledge. This requires a “dialogue of ways of
knowing” wherein theoretical-technical results and local cultural
creativity and intuition come together to produce new strategies and
technologies for ecological restoration (Leff, 2010). Especially at large
spatial scales, effective strategies to restoration in drylands should be
based on integrated biophysical and socioeconomic evaluation methods
(Costantini et al., 2016). As is well known, these are areas that still need
strong development in relation to the science, business, economy, pol-
itics, and practice of ecological restoration. We may take as a model to
emulate the work of Elliot and co-workers over many years in their
application and development of the FSA in northern Thailand. They not
only addressed obstacles in methods for growing in nurseries (Elliott
et al., 2003), but also recognized the need to work in collaboration with
local populations of farmers who depend on traditional knowledge of
plants and of prevailing climatic and environmental conditions. Thus,
Elliot and co-workers include local villagers in the definition of objec-
tives, choice of framework species, search for seed sources, and the task
of producing plants in nurseries is indeed exemplary (Elliot and
Kuaraska, 2008). In the recent book-length contribution, Elliot et al.
(2013) also describe in detail the obstacles to be overcome with respect
to scaling up of this participative approach. Examples include the need
D.R. Pérez et al.
Fig. 3. (A): Survival percentages 11 months after plantation in the five planting sites (S1-S5) of AL: Atriplex lampa, HA: Hyalis argentea var. latisquama, and SS: Senecio
subulatus var. subulatus. (B): Survival rates 22 months after plantation of AL: A. lampa, HA: H. argentea var. latisquama and SS: S. subulatus var. subulatus. Bar groups
that do not share a common letter differ significantly (p < 0.05).
for training and education with regards seed collection and methods for
growing in nurseries.
4.1.3. Concern 3: different spatial layout and mechanically engineered
microsites
The spatial distribution of candidate dryland framework species in
plantations is an issue that should be evaluated and tested in the future.
Initially it is necessary to evaluate how the woody plant species under
consideration for use in a FDSA effort differ in their capacity to facil-
itate the recruitment of additional native species without direct inter-
vention. This possibility is likely to vary according to the competition
that can be generated by the root systems and associated pathogens or
mutualists, the microclimate that is produced by the canopy of the
various species, and eventually, in some species, the production of re-
sins or allelopathic substances (Pugnaire, 2010; González and Pérez,
2017). Additionally, the effectiveness and longevity of varying ar-
rangements like nucleation based on island planting (Hulvey et al.,
2017), or other types of engineering and design should be field-tested in
dryland restoration work. Additionally, an option to be considered, on
7
Journal of Arid Environments 161 (2019) 1–10
the basis of both theoretical and empirical considerations, is the judi-
cious use of mechanical barriers in plantation sites (Li et al., 2006). The
placing of permanent or temporary mechanical barriers perpendicular
to dominant winds and/or water flow, can potentially reduce the loss of
resources (soil, water, and seeds) and reduce evapotranspiration (Fick
et al., 2016). When grown successfully, framework species can also
offer biological barriers to wind that in turn increase site attractiveness
for native fauna. As noted above, complementary techniques to increase
effectiveness in the spontaneous recruitment and establishment of na-
tive plants at sites distant from planting sites may also need to be
considered with both conceptual and systems modelling (Noy-Meir,
1973) as well as experimental approaches (James et al., 2013). An
additional intervention to test is manipulation and remodeling of soil
surfaces by pitting and other means to increase rain-on microsites. The
goal here is to enhance the usefulness of seed rain and seeds moved by
animal dispersers, and thus to promote the spread of plant cover, and
the progressive increase of new habitats and foci for spontaneous re-
covery (DeFalco et al., 2012).
D.R. Pérez et al. Journal of Arid Environments 161 (2019) 1–10

Fig. 4. (A): Plant height at planting and 11 months after plantation. (B): Shoot biomass at planting and 11 months after plantation. (C): Root biomass at planting and
11 months after plantation. D): Comparison between the crown area of seedlings planted and plants of the reference ecosystem (ER) and the planting sites (S). HA:
Hyalis argentea var. latisquama, AL: Atriplex lampa and SS: Senecio subulatus var. subulatus. Different letters a-b/'-b'/a''-b'' indicate significant differences (p < 0.05).

4.1.4. Concern 4: effective monitoring from drones for adaptative
management
Importantly, images that can be obtained quickly and with a great
level of detail by unmanned aerial vehicles (drones), equipped with
various imaging technologies could be used to rapidly and efficiently
monitor and help evaluate the survival and growth of out-planted
seedlings/saplings rapidly, over large areas. Such technologies are
especially useful in drylands, since vegetation coverage is generally
low, with few strata, and it may be possible to use machine-learning to
develop automated species recognition systems capable of counting
individuals of each plant species reintroduced artificially and
estimating plant cover of each species that survives over time.
Similarly, rapidly advancing aerial imagery technology and software
should allow monitoring of the arrival of new species that establish
themselves spontaneously, either under nurse plants in the plantations
or colonization of bare soil sites.
4.2. Attributes for DFSA
4.2.1. Attribute a: survival and growth
Achieving high rates of survival in the first year after outplanting is
critical to increasing revegetation and restoration effectiveness (Abella

Table 3
Faunal groups attracted by and plant parts eaten for Atriplex lampa (AL), Senecio subulatus var. subulatus (SS), and Hyalis argentea var. latisquama (HA).(1): Reference
for A. lampa (Zarco, 2016). (2): References for A. lampa and H. argentea var. latisquama (Puig et al., 2006). (3): References for A. lampa (Borruel et al., 1998; Puig
et al., 2006) and H. argentea var. latisquama (Puig et al., 2006).*: Information provided by rural residents.
Assessed fauna group

Stems/leaves Fruits Nests/burrows Resting/feeding/hunting

(1) (1)
Granivorous birds Confirmed (AL-SS) Confirmed (AL-SS-HA) Probable (AL-SS-HA) Observed (AL-SS)(1)
Large wild herbivores or omnivores Confirmed (AL-HA)(2) Confirmed (AL-HA)(2) (SS) No No
Medium wild herbivores or omnivores Probable (AL-HA)(3) Confirmed Confirmed No
(AL-SS-HA) (AL-SS-HA)
Small wild herbivores Probable (AL-HA)(3) No Confirmed No
(AL-SS-HA)
Domestic herbivores Confirmed (AL-HA)(2) (SS) Confirmed (AL-HA)(2) (SS) No No

8
D.R. Pérez et al.
et al., 2012; Yirdaw et al., 2017). In humid tropical and seasonally dry
tropical forests, the tolerance of species to open spaces with no shade is
a central criterion. Given the recurrent difficulties encountered in fa-
cilitating the initial establishment phase of plants planted or sown in
drylands, survival and growth rate qualify as the first key attribute to
consider in the selection of species for DFSA. To wit, both A. lampa and
H. argentea maintained the same rates of survival 11 and 22 months
after plantation; in contrast, S. subulatus showed high mortality in the
second year. This was apparently not linked to herbivory or climatic
factors since no evidence of attack by herbivores was observed and
mortality was variable only with respect to varying soil conditions (see
soil electrical conductivity variability of the sites in Table 1 and the
correlation with infiltration in the results). The results suggest the need
to evaluate species and possible groups of different framework species
for each type of soil (e.g., salinized, compacted, aeolian accumulations,
and desert pavement). The mortality of S. subulatus var. subulatus, a
species typically found growing in hydraulically eroded soils, dunes,
and alluvial fans (Fernandez et al., 2015) in the second year, also shows
us that not all species of the reference ecosystem in drylands can adapt
to any type of disturbance or change in soil properties and therefore for
DFSA the most adaptable species should be evaluated. The results
presented here show that 22 months after out-planting the crown area
of all three species showed rapid growth, which is essential for suc-
cessful application of DFSA. In drylands, where soil erosion is often a
great obstacle, we should consider underground and rhizomatous
growth potential of candidate species. Among our candidate species, as
mentioned already, vigorous rhizomatous growth of Hyalis argentea var.
latisquama enables it to spread horizontally, and therefore help retain
and consolidate degraded soils.
4.2.2. Attribute b: attractiveness for fauna
Relative attractiveness to and interaction with native animal and
insect dispersers should be evaluated in terms of palatability, refuge
habitat, and nesting niches offered to potential dispersers in drylands.
Fauna that are abundant in drylands need shade for thermoregulation
such as can be found in plantations and the animals can in turn con-
tribute to bioperturbation and in some cases seed and fruit transport.
Perturbation by mammals for example can be an important driver in
pedogenesis, in structuring landscapes, and in maintaining hetero-
geneity in arid ecosystems (Whitford and Fenton, 1999). Ultimately,
native fauna will play a crucial role in the creation, linkage, and ex-
pansion of new and pre-existing vegetation far from the initial planta-
tion sites designed and established with the help of DFSA. Additionally,
possible disadvantages of certain plant-animal interactions should also
be considered in some cases, in the case of candidate framework species
being particularly attractive to domestic livestock or damaging invasive
fauna. In the future, with more species evaluated as candidates for use
within a DFSA, and more studies taking advantage of photographic
traps, drones, meta-barcoding, and other tools and techniques, it is
possible that some animal species are found to have particular ad-
vantages for restoration work. No indication of this factor showed up in
the bibliographic study we carried out for our study area.
4.2.3. Attribute c: easy to germinate
Abundant seeds and are easy to grow are ideal for restoration work
and for use as framework species (Goosem and Tucker, 2013) and for
DFSA. It is necessary to progressively look for treatments to promote
seed germination that are inexpensive and simple to apply, even by
people with no professional training or access to lab facilities
(Rodriguez Araujo et al., 2017). This is the case of S. subulatus var.
subulatus; H. argentea var. latisquama, and A. lampa, our three local
species with rapid and easy germination.
At the same time, it will be necessary in the future, to study the
autecology and reproductive phenology of candidate species to identify
those that most quickly produce viable seeds. Our observations indicate
that already in the second spring post-plantation, both S. subulatus and
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Journal of Arid Environments 161 (2019) 1–10
H. argentea var. latisquama begin to produce flowers, whereas A. lampa
in ours plantations only blooms beginning three to five years post-
plantation. This is a key topic to address in order move forward with the
development and application of our DFSA.
5. Conclusions
Our results show that although we face a very demanding set of
challenges both in research and restoration practice, it is possible to
identity some subset of native species in dryland ecosystems with po-
tential to help kickstart reintroduction, revegetation, and other ecolo-
gical succession processes essential to effective ERR. These criteria can
then be integrated into a methodology that we call DFSA. This approach
can be integrated with complementary sets of selection criteria in a
given project or program, and with pre-existing models and techniques,
e.g., state and transition modelling, landscape functional analysis,
spatial analysis, and predictive systems models that capture the prob-
abilistic nature of ecosystem response to management practices and
ERR interventions (James et al., 2013; Carretero and Dalmasso, 2015;
Costantini et al., 2016). In summary, future lines of research and action
for the use of FSA in drylands, especially for medium- and large-scale
restoration and rehabilitation projects include:
a Ongoing exploration in the varying habitats of the ecosystem of
reference for species or ecotypes capable of surviving and thriving
on different soil types, despite drought and other harsh conditions.
This should be carried out in conjunction with the development of
easy and effective and low-cost propagation and planting techniques
for the candidate species. These species should not only be easy to
reproduce, but also able to grow rapidly. The search for and use of
micro-symbionts to use with each candidate plant species is another
important topic for research and development.
b Protocols to evaluate candidate DFSA species from varying habitats
in the ecosystem of reference, and to study the autecology and re-
productive phenology of candidate species to identify those that
most quickly produce viable seeds.
c Greater empowerment and inclusion of local human communities is
needed in all aspects of restoration activities. This implies building a
plural and closed dialogue of knowledge between local and in-
digenous communities. In this way, it is possible to improve the
selection and evaluation of the species, control the physical, human,
intellectual and financial resources, and foster the capacity to work
together to identify needs and problems, collect information, make
collective decisions and implement efforts. interventions in a mu-
tually agreed manner
d As suggested above, the use of drones can greatly increase the ca-
pacity to analyze progress of plantations and the advent of un-
planted species on medium- and large spatial scales. Indeed, in a
subsequent contribution, we will report on our own on-going re-
search with drones.
e With this information, and multiple field applications with careful
monitoring and evaluation across a range of dryland contexts, we
suggest that a Dryland Framework Species Approach can developed
further and emerge as a valuable conceptual and methodological
tool to guide the development, monitoring, and application of eco-
logical restoration and rehabilitation activities in drylands. We
particularly call for the establishment of long-term studies carried
out on institutionally and financially stable sites where all con-
ceptual, socio-economic, and practical aspects of DFSA-oriented
restoration and rehabilitation can be field tested and improved in
conjunction with local communities and other major stakeholders.
Acknowledgements
We are grateful to Juana Lagos for her help with field work and to
Guillermo Sabino for assistance in statistical analysis. We also thank
D.R. Pérez et al. Journal of Arid Environments 161 (2019) 1–10

Javier Contreras and Joaquín Pérez Carrió, the gardeners in charge in (Eds.), Restauración ecológica en la diagonal árida de la Argentina. Vazquez Mazzini
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Goosem, S.P., Tucker, N.I.J., 1995. Repairing the Rainforest: Theory and Practice of
plantations were funded by Project 04/U014 and by the technical col- Rainforest Re-establishment in North Queensland's Wet Tropics. Wet Tropics
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