Agriculture, Ecosystems and Environment 289 (2020) 106736

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Agriculture, Ecosystems and Environment

journal homepage: www.elsevier.com/locate/agee

Predicting how seed-eating passerines respond to cattle grazing in a semi- T

arid grassland using seed preferences and diet
M. Cecilia Sagarioa, Víctor R. Cuetob, Agustín Zarcoc,d, Rodrigo Polc,d, Luis Maronec,d,*
a
Ecodes, Instituto de Investigaciones en Biodiversidad y Medioambiente (INIBIOMA – CONICET and UNComahue), and Centro de Ecología Aplicada de Neuquén (CEAN),
Junín de los Andes, Neuquén, Argentina
b
Ecodes, Centro de Investigación Esquel de Montaña y Estepa Patagónica (CIEMEP – CONICET and Facultad de Ciencias Naturales, UNPatagonia), Esquel, Chubut,
Argentina
c
Ecodes, Instituto Argentino de Investigaciones de las Zonas Áridas (IADIZA – CONICET), Mendoza, Mendoza, Argentina
d
Facultad de Ciencias Exactas y Naturales, UNCuyo, Mendoza, Mendoza, Argentina

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

Keywords: Numerical responses of animals to habitat perturbation often seem inconsistent, spreading skepticism about the
Bird abundance predictive capacity of applied ecology. Domestic grazing changes several habitat variables that can affect seed-
Feeding flexibility eating birds. Birds, in turn, show adaptations (e.g. in their feeding behavior) that could allow them to overcome
Forb seeds habitat perturbations. Here we modelled habitat variables (e.g. cover of different plants, panicles, soil seed bank)
Grass seeds
in grazed and ungrazed (or lightly grazed) habitats of the central Monte desert, Argentina, to detect those
Grazing of semi-arid grasslands
affected by grazing activity. There was no effect of grazing on shrub and tree cover, but grazing reduced the
Predictive ecology
abundance mostly of large grass seeds but also of small grass and forb seeds. Then, we used model’s outputs and
knowledge of feeding preferences of the five most common seed-eating passerines in the Monte to make species-
specific predictions: changes in abundance of grass seed specialists (Saltatricula multicolor, Microspingus torquatus
and Porphyrospiza carbonaria) due to grazing activity should be consistent and should depend on large grass
seeds (i.e. preferred seeds), whereas changes in abundance of more generalist species (Zonotrichia capensis and
Diuca diuca) should be less consistent and explained also by the abundance of other seeds. The abundance of
large grass seeds was sufficient to predict the abundances of S. multicolor, M. torquatus and P. carbonaria. The best
model for predicting the abundance of Z. capensis included large grass seeds as well as small grass or forb seeds.
No model including the abundance of seeds predicted the abundance of D. diuca. Therefore, feeding behavior
explained the abundance of four out of the five bird species. A review of the literature showed that feeding
behavior is also a good predictor of habitat use in other desert grasslands. Conservative range management
should consider, and even manipulate, the level of the seeds preferred by wildlife. Grazed grasslands should be
rested from grazing on a rotational basis so that grasses, especially those whose seeds are preferred by birds, can
seed.

1. Introduction
Land use changes and habitat disturbance are major drivers of ve-
getation change and biodiversity loss (Díaz et al., 2007). Livestock
grazing is the most globally widespread land use, encompassing 25% of
Earth’s land surface, and it is even more pervasive in arid and semiarid
ecosystems (Frank et al., 2013), where it usually differs from the nat-
ural grazing regime. A key issue for the conservation and management
of desert birds is to determine which species are negatively or positively
affected by grazing and why such opposite responses occur. Differences
in ecological context, such as climate (Bock and Bock, 1999), the height
of natural grasslands (Isacch and Cardoni, 2011), or the stocking rates
(Isacch and Cardoni, 2011), as well as differences in bird traits like
feeding behavior (Martin and Possingham, 2005; Seymour and Dean,
2010; Camín et al., 2015), have been correlated with, or invoked to
explain, the responses of bird species to grazing.
In the ecological literature, such responses are sometimes incon-
sistent, and researchers frequently associate inconsistent results with
the contingent nature of animal behavior (Fox and Morrow, 1981;
Gordon, 2011), spreading skepticism on the predictive capacity of their
discipline. Nevertheless, the pervasiveness of contingent behavior
should be empirically tested instead of assumed (Gordon, 2011; Marone

⁎
Corresponding author at: Ecodes, Instituto Argentino de Investigaciones de las Zonas Áridas (IADIZA – CONICET), CC 507, 5500, Mendoza, Argentina.
E-mail address: lmarone@mendoza-conicet.gob.ar (L. Marone).

https://doi.org/10.1016/j.agee.2019.106736
Received 3 September 2019; Received in revised form 25 October 2019; Accepted 26 October 2019
0167-8809/ © 2019 Elsevier B.V. All rights reserved.
M.C. Sagario, et al. Agriculture, Ecosystems and Environment 289 (2020) 106736

Fig. 1. Map showing localities where studies were carried out: Telteca Natural Reserve (T), Biosphere Reserve of Ñacuñán (Ñ), and General Alvear (A), in the Monte
desert of Argentina (black area). High-resolution Google Earth imagery indicates the study sites subjected to none or low (A) and high (B and C) grazing pressure in
each locality. Black lines indicate the boundary of Telteca and Ñacuñán reserves.

et al., 2015), because inconsistent results do not only depend on nature
variability but also on the research approach employed (e.g. sampling
design and correlations). Through classical null-hypothesis testing, re-
searchers have reported no significant effects (e.g. Medin, 1986; Davis
et al., 1999), negative effects (Bock and Bock, 1999), positive effects
(Powell, 2008), or any mixture of negative, positive and null effects
(Grzybowski, 1983; Bock et al., 1984; Gordon, 2000; Gonnet, 2001;
Fontaine et al., 2004) of cattle grazing on the abundance of seed-eating
birds, sometimes for the same habitat and species (Martin and
Possingham, 2005). Although opposing results can arise from the
variability of the multiple factors involved in domestic grazing (e.g.
stocking density, grazing history, habitat type and season assessed),
they are disappointing because ecologists look for repeatable and con-
sistent patterns. A different though complementary approach is un-
veiling the mechanisms that act as causal links between habitat al-
teration and niche bird properties to produce a priori hypotheses on the
responses of bird species to grazing (Martin and Possingham, 2005;
Marone et al., 2008; Kutt and Martin, 2010; Seymour and Dean, 2010;

2
M.C. Sagario, et al.
Pol et al., 2014; Ehlers-Smith et al., 2015). This tactic relies on the
assumption that birds have evolved within the pressures imposed by
their environments (e.g. food availability) and that some bird traits are
therefore adaptations (e.g. feeding flexibility), which can be used to
elaborate species-specific predictions (Seymour and Dean, 2010).
Subtle differences in the feeding behavior of seed-eating birds
wintering in the central Monte desert, Argentina, offer an opportunity
to build such species-specific predictions. Regarding the composition of
the granivorous field diet in natural undisturbed habitats, Saltatricula
multicolor (Many-colored Chaco Finch), Microspingus torquatus (Ringed
Warbling-Finch), and Porphyrospiza carbonaria (Carbonated Sierra-
Finch) eat and select (i.e. they consume items in higher proportions
than expected according to their availability) grass seeds almost ex-
clusively, especially large grass seeds, whereas Zonotrichia capensis
(Rufous-collared Sparrow) and Diuca diuca (Common Diuca-Finch) have
more mixed diets, with grass as well as forb seeds (Lopez de Casenave
et al., 2008; Marone et al., 2008; Sánchez and Blendinger, 2014). In
grazed areas, S. multicolor and M. torquatus continue to feed mostly on
large grass seeds, but Z. capensis and D. diuca are capable of moderate
diet switching, incorporating a higher proportion of small grass seeds or
forb seeds (Marone et al., 2017). Concerning seed preferences, estab-
lished through laboratory experiments in which seeds of different spe-
cies are offered simultaneously and with equal availability, S. multicolor
preferred 65% of the grass species offered and 0% of the forb species
offered, D. diuca preferred 65% of the grass species offered and 25% of
the forb species offered, and Z. capensis preferred 90% of the grass
species offered and 40% of the forb species offered (Camín et al., 2015).
In summary, S. multicolor, M. torquatus, and P. carbonaria seem to be
specialized grass seed foragers that show even less feeding flexibility in
the field than that expressed in the preference trials (Cueto et al., 2006).
By contrast, Z. capensis and D. diuca would be more flexible foragers
that could incorporate less preferred seeds when preferred ones are
scarce in the habitat (Camín et al., 2015; Marone et al., 2017). We
assume that in relatively simple environments like desert grasslands,
detailed knowledge of the feeding behavior of animals may be sufficient
to predict the reciprocal effects between them and their food resources,
and hypothesize that (a) accordingly to plant preferences of domestic
grazers (Pol et al., 2014), they reduce the cover and provoke changes in
the composition of the grass layer, decreasing grass seed availability
(specially of large grass seeds), but grazers have lower effect on forb
seeds, and (b) knowledge of feeding behavior of birds (and of their
resources) is efficient to predict bird numerical responses to domestic
grazing. Therefore, under grazing pressure, the reduction in the abun-
dance of the strictly grass-seed specialist birds (S. multicolor, M. tor-
quatus, and P. carbonaria) should be consistent and explained by the
reduction of large grass seeds. By contrast, numerical reductions of the
more flexible and generalist bird species (Z. capensis and D. diuca)
should be inconsistent or even absent, because their abundance may
depend on large grass seeds but also on other seeds that may be less
affected by grazing.
2. Materials and methods
2.1. Study area and design
The study was carried out in open woodlands of Prosopis flexuosa in
three localities of the central Monte desert, Argentina: Telteca (32° 02′
S, 68° 00′ W), Ñacuñán (34° 03′ S, 67° 54′ W), and General Alvear
(hereafter Alvear, 35° 06′ S, 66° 40′ W). These localities are on the
plains at the eastern foothills of the Andes (Fig. 1). The region is arid
and seasonal, with hot, humid summers and dry, cold winters. Annual
rainfall is heavily concentrated (approximately 75%) in spring and
summer. Climate in Telteca is drier and warmer than in the other two
localities, but habitat structure and main plant species are very similar
for all three localities (Table 1). From mid 1900s on, the main economic
activity in all localities has been continuous cattle grazing (mostly cows
3
Agriculture, Ecosystems and Environment 289 (2020) 106736
and, to a lesser extent, horses in Ñacuñán and Alvear; and goats in
Telteca), and the extraction of firewood from shrubs and trees. The area
is currently free of large native wild grazers.
We established one or two sites with high domestic grazing pressure
and one with none / low domestic grazing pressure in each locality.
Selected sites were surrounded by areas of over 1000 ha with similar
grazing pressure. Within the Telteca Reserve, one site had been under
low grazing pressure since 1986 (TLG), but the other had been sub-
jected to the more intense grazing that is typical of the region (THG). In
Ñacuñán, one site was located inside the Reserve of Ñacuñán where
cattle have been excluded for > 40 years (ÑLG), and sites ÑHG1 and
ÑHG2 were located on neighboring cattle ranches. In Alvear, both sites
were located on cattle ranches but one of them (ALG) had been exposed
to grazing to a lesser extent than the other (AHG). All study sites
were > 1.5 km apart from each other (see Fig. 1), and their grazing
regime remained very similar during the study period. The soil and
general appearance of vegetation in contrasting sites was similar for all
localities (see 3. Results).
2.2. Grazing pressure
There are no reliable records of the actual livestock density in the
study areas, but the presence of dung is an indicator of the overall
grazing pressure that may reflect both current and recent livestock
activity (Cingolani et al., 2002). In July 2013, we recorded the occur-
rence of dung of cows, horses and goats regardless of its degree of de-
composition. In each locality, we sampled in 16 transects 300 m long,
located randomly at the paired sites representing each grazing condi-
tion, except for ÑHG1 and ÑHG2 with n = 8 transects each. In every
transect we recorded the presence or absence of dung in a 50 cm
quadrat sampler which was located every 10 m at a random distance
between 0 and 5 m from each side of the transect line. We used the
proportion of quadrats with dung present at each site in every location
as a proxy for grazing pressure (von Müller et al., 2012). Results of
these samplings for Ñacuñán and Telteca have partially been published
in Pol et al. (2014).
2.3. Vegetation and soil seed bank
We interspersed 20–40 parallel transects (25 m long, > 20 m apart)
at every site to measure plant cover and the number of panicles.
Vegetation samplings were performed in March-May, at the end of the
growing and reproductive season of most perennial grasses, in 2009 in
Telteca (THG n = 37, TLG n = 39), 2010 in Ñacuñán (ÑHG1 n = 20,
ÑHG2 n = 20, ÑLG n = 20), and 2012 in Alvear (AHG n = 20, ALG n
= 20). We recorded plants to the level of species or genus that touched
a vertical pole at 25 random sampling points within each transect.
When the plant was an adult individual of grass (i.e. plants producing
new seeds or that had previously produced seeds) we also recorded the
number of panicles. We calculated the percentage cover of grasses
(number of touches of plant X / total number of touches), forbs, low
shrubs (< 1 m), tall shrubs (> 1 m), and trees, and the number of grass
panicles for each transect.
We also evaluated the composition and size of the soil seed bank at
every site during October-November 2011–2014 in Telteca, Ñacuñán,
and Alvear. In each year and site, we collected 120 soil samples, except
for sites in Ñacuñán where n = 60 (ÑHG1, ÑHG2, and ÑLG). We ran-
domly allocated sampling points for soil core samples according to the
cover of the main microhabitats in the landscape: beneath the tree
canopy (15% in both grazing conditions) and the tall shrub canopy
(35% in both grazing conditions), under low shrubs (13% in both
grazing conditions) and grasses (17% and 7% in ungrazed and grazed
sites, respectively), and on bare soil (20% and 30% in ungrazed and
grazed sites, respectively). We collected soil samples using a cylindrical
sampler, 3.2 cm in diameter and 2 cm deep (80% of seeds are found in
the upper 2 cm of soil). In the laboratory, soil samples were sifted
M.C. Sagario, et al. Agriculture, Ecosystems and Environment 289 (2020) 106736

Table 1
Main climatic and vegetation features for the three localities in the central Monte desert, Mendoza province, Argentina, where we conducted the study. Both historic
mean annual rainfall and mean annual rainfall during bird sampling (2013–2015, between brackets) are included.
Telteca Ñacuñán Alvear

Mean annual rainfall 155 mm (184 mm) 334 mm (310 mm) 329 mm (317 mm)
Mean annual temperature 18.7 °C 15.6 °C 15.4 °C
Main trees Prosopis flexuosa (Fabaceae) P. flexuosa P. flexuosa
Geoffroea decorticans (Fabaceae) G. decorticans G. decorticans
Main shrubs Larrea divaricata (Zigophyllaceae) L. divaricata L. divaricata
Tricomaria usillo (Malpighiaceae) Capparis atamisquea (Capparaceae) Larrea cuneifolia (Zigophyllaceae)
Bulnesia retama (Zigophyllaceae) Condalia microphylla (Rhamnaceae) C. microphylla
Lycium spp. (Solanaceae) Atriplex lampa (Amaranthaceae)
Acantholippia seriphioides (Verbenaceae) Prosopis alpataco (Fabaceae)
Main grasses Perennials Perennials Perennials
Trichloris crinita T. crinita T. crinita
Aristida mendocina A. mendocina A. mendocina
Setaria spp. Setaria spp. Panicum urvilleanum
Pappophorum spp. Pappophorum spp.
Sporobolus rigens Sporobolus cryptandrus
Annuals Digitaria californica
Bouteloua aristidoides
B. barbata
Aristida adscensionis
Source González Loyarte et al. (2000) Lopez de Casenave (2001) Cesca et al. (2014)
This study This study This study

through a 0.27 mm mesh sieve (the smallest seeds recorded in the ha-
bitats did not pass the sieve) and then washed in the same sieve under
water pressure and air-dried. We searched for sound seeds under a
stereoscopic microscope and identified seeds to the species or genus
level using a reference collection. Seeds recorded were ascribed to one
of the three plant groups that are systematically consumed by birds:
small grass seeds (< 0.15 mg), large grass seeds (0.16 – 0.90 mg), and
forb seeds of all sizes (Marone et al., 2008). Results of these samplings
for Ñacuñán and Telteca have partially been published in Pol et al.
(2014).
2.4. Birds
We studied the five most abundant granivorous species present in
the three localities in winter: M. torquatus, S. multicolor, D. diuca, P.
carbonaria (all Thraupidae), and Z. capensis (Emberizidae). All species
are present throughout the year in the three localities (del Hoyo et al.,
2011). Though, except for S. multicolor, abundances are usually higher
during the winter, and some evidence suggests that at least some por-
tions of the populations are migratory (Cueto et al., 2011; del Hoyo
et al., 2011; Sagario et al., 2014). We surveyed bird abundance in the
austral winter (mid July-mid August) during three years (2013–2015).
Sampling in both grazing conditions was conducted during two to four
consecutive days at each locality. We estimated bird abundance using
strip transects, where the recorded number of individuals of every
species is divided by the area covered by the strip transect (Emlen,
1977). Trailside strips were narrow enough to be able to detect all bird
cues (e.g. songs, calls, and sightings). The strip width was of 25 m on
each side of the transect for every species. We counted birds along
300 m long transects (n = 8 at each site, scattered within an approx-
imate area of 100–150 ha), separated a minimum of 250 m to avoid
double counts and reduce spatial dependency. Transects were located at
least 200 m from roads or external fences to avoid edge effects. Each
transect survey was performed by a single observer, and only two ob-
servers (MCS and AZ) performed surveys during the entire study. Ob-
servers and the order of transects were alternated to avoid bias. Surveys
were not conducted during periods of rain or strong winds. We recorded
the presence of individuals during the first five hours after sunrise
(Verner, 1985). We did not count any birds that flew across the strip
transect. Each transect was surveyed three times in every sampling
period.
2.5. Statistical analyses
We tested the effect of grazing on dung presence, vegetation, and
soil seed banks using generalized linear mixed models (GLMMs). We
modeled data using Poisson (number of large and small grass seeds per
sample), Negative Binomial (number of forb seeds per sample and
number of panicles per transect), and Binomial distributions (propor-
tion of dung per transect, and cover per transect of grasses, low shrubs,
tall shrubs, and trees), considering the temporal (year) and/or spatial
(locality) non-independence of sites. The most complete of these models
included grazing as a fixed effect, and a random slope (grazing effect)
and intercept (locality or year*locality depending on the sampling de-
sign) as random factors. We used backwards model selection utilizing
Akaike Information Criteria and log-likelihood tests to choose for the
best random structure and then estimated parameters for the best
model (Zuur et al., 2009).
We tested whether mean abundance of large grass seeds, small grass
seeds and/or forb seeds in each site predicted bird counts using GLMMs.
The number of small grass seeds and forb seeds was highly correlated
(r = 0.981, p < 0.001) so we included only one type of these seeds
(small grass seeds) in each model to avoid problems with multi-
collinearity. Main results did not change when including forbs instead
of small grass seeds in the models. We modeled total bird counts using
Poisson or Negative Binomial distributions considering the spatial (lo-
cality) non-independence of sites. Total bird count for each transect was
calculated as the sum of all counts made on that transect (i.e. the sum of
nine counts: three counts per year during three years). Levels of the
fixed effects (i.e. the number of different types of seeds) were the mean
number of seeds (2011–2014) for each site. The most complete model
included the different type of seeds as fixed effects, and locality as a
random factor. Each bird species was modeled in separate GLMMs. For
every species, we used backwards model selection utilizing Akaike
Information Criteria and log-likelihood tests to choose for the best
random structure and for the best fixed structure and, finally, we esti-
mated parameters for the best model (Zuur et al., 2009).
All statistics were conducted in R (R Core Team, 2016), and GLMMs
were implemented in the ‘lme4’ (Bates et al., 2015) or ‘MASS’ (Venables
and Ripley, 2002) packages.
3. Results
We did not find dung in any transect of the ungrazed site in

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M.C. Sagario, et al. Agriculture, Ecosystems and Environment 289 (2020) 106736

Fig. 2. Dung proportion, plant cover by functional group,

number of panicles per plant of perennial grass, and of seeds
of the three groups usually consumed by seed-eating birds in
the soil bank, in sites under none or low (black bars) and high
(white bars) grazing pressure in Prosopis open woodlands at
three localities (Telteca, Ñacuñán, and Alvear) in Mendoza
province, Argentina. The values (mean + SE) come from the
average of the average of the variables in each locality (n = 3
for each bar of the graph) for each type of site. Asterisks in-
dicate a significant effect of grazing detected using GLMM
with grazing as a fixed effect and grazing, locality and/or year
as random factors (see Table 2 for detailed models, **
p < 0.01, *** p < 0.001). Note that vertical axes differ in
scale.

Table 2
Estimates for the best generalized linear models (GLM) or generalized linear mixed models (GLMM) describing grazing effect on seeds (large and small grass seeds
and forb seeds, n = 120 for every grazing condition in each locality and year), vegetation (grass cover, number of panicles and cover of low shrubs, tall shrubs, and
trees, n = 20–30 for every grazing condition in each locality) and a proxy for grazing pressure (dung presence, n = 16 for every grazing condition in each locality)
recorded in sites under none or low and high grazing pressure at three localities (Telteca, Ñacuñán and Alvear) of the Monte desert, Mendoza, Argentina. P-values and
coefficients for significant effects of grazing are in bold.
Grazing (fixed effect) Random Effects

Response variable Int. Coef. Lower 95% CL Upper 95% CL Z p-value Slope | Intercept SD

Dung* −0.67 0.98 0.79 1.16 10.31 < 0.001 | locality 0.13
Low shrubs −1.92 0.73 −0.16 1.61 1.62 0.106 Grazing | locality 0.71 | 0.76
Tall shrubs −0.76 −0.32 −0.78 0.14 −1.38 0.167 Grazing | locality 0.25 | 0.38
Trees −1.64 0.13 −0.05 0.31 1.43 0.151 – –
Grasses −0.59 −1.03 −1.54 −0.52 −3.95 < 0.001 Grazing | locality 0.22 | 0.42
Panicles 4.32 −2.34 −2.95 −1.73 −7.51 < 0.001 | locality 0.85
Large grass seeds −0.33 −1.44 −1.77 −1.11 −8.46 < 0.001 Grazing | Year*locality 0.49 | 0.52
Small grass seeds −1.54 −0.95 −1.62 −0.28 −2.79 0.005 Grazing | Year*locality 1.91 | 0.85
Forb seeds 0.43 −0.60 −0.98 −0.21 −3.03 0.002 Grazing | Year*locality 1.45 | 0.61

* This model only includes two localities (Telteca and Alvear). We excluded Ñacuñán because of the total lack of dung in the site under nil grazing.

Ñacuñán, so we excluded this locality from the statistical analysis since
the comparison was trivial (dung proportion was 0.31 ± 0.02 in
ÑHG1, and 0.46 ± 0.04 in ÑHG2). As expected to confirm a good
choice of sampling sites (i.e. sites within each locality differing in
grazing activity), the overall average proportion of dung was highest at
sites under high grazing (Fig. 2), and the model including Telteca and
Alvear showed that grazing was associated with an increase of 71% in
the presence of dung (Table 2). Regarding woody cover, the greatest
random variation among localities occurred in the cover of low shrubs
(Table 2), but the overall results showed that there was no grazing ef-
fect neither in the cover of these shrubs nor in the cover of high shrubs
or trees (Fig. 2, Table 2). The effect of grazing on the seeds most con-
sumed by birds varied between localities and years (Fig. 3, Table 2), but
its overall effect was negative on the three types of seeds. The largest
reduction was observed for large grass seeds (76%), followed by small
grass seeds (62%), and forb seeds (45%, Fig. 2, Table 2). Other aspects
related to the abundance of grass seeds were also reduced by grazing:
grass cover (53%), and the number of panicles (90%; Fig. 2, Table 2).
The abundance of birds in sites under high and low grazing condi-
tions varied over the years and localities but was usually higher in sites
with low grazing (Fig. 4). The models showed that in four of the five
species changes in abundance were related to the abundance of seeds in

5
M.C. Sagario, et al. Agriculture, Ecosystems and Environment 289 (2020) 106736

Fig. 3. Mean abundance + SE of the three

group of seeds usually consumed by seed-
eating birds in the soil seed bank during con-
secutive years in sites under none or low (black
bars) and high (white bars) grazing pressure in
three localities of the central Monte desert,
Argentina. The values come from the average
for each locality and grazing condition
(n = 120 for each bar of the graph, except for
the low grazing site in Ñacuñán where n = 60).
Note that vertical axes differ in scale.

Fig. 4. Mean abundance + SE of birds during consecutive years in sites under none or low (black bars) and high (white bars) grazing pressure in three localities of the
central Monte desert, Argentina. The values come from the average (three years) of the average (three counts per year) for each transect and grazing condition (n = 8
for each bar of the graph, except for high grazing sites in Ñacuñán where n = 16). Note that vertical axes differ in scale.

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M.C. Sagario, et al. Agriculture, Ecosystems and Environment 289 (2020) 106736

Table 3 abundance of four out of the five seed-eating bird species studied, and
Model selection for the effect of different types of seeds on counts of five species the species-specific feeding behavior of those birds explained such nu-
of birds in three localities (Telteca, Ñacuñán, and Alvear) of the Monte Desert, merical patterns.
Mendoza, Argentina. Large: Large grass seeds. Small: Small grass seeds. The Although given its general behavior Z. capensis may be considered a
abundance of small grass seeds was highly correlated with the abundance of
generalist species (Ríos et al., 2012; Camín et al., 2016), its feeding
forb seeds (r = 0.981, p < 0.001), so we only included the abundance of small
behavior is more restricted. Z. capensis consumes seeds of different fa-
grass seeds in the models. Best models were the same when including forb seeds
milies but predominantly prefers grass seeds (Cueto et al., 2001; Camín
instead of small grass seeds. RE: Random effect (locality). Best models are in
bold. We used Poisson distribution for Saltatricula multicolor and Microspingus et al., 2015), and should be better classified as an “expanding specia-
torquatus (the latter was corrected for overdispersion using Quasipoisson), and list” (Heller, 1980), i.e. a species that feed on grass seeds until they fall
Negative Binomial distribution for Zonotrichia capensis, Porphyrospiza carbonaria below some threshold value at which the bird begins to include less
and Diuca diuca. preferred or alternative food types (e.g. forb seeds). For example, Z.
capensis moderately switchs its diet in grazed areas by including a larger
Species Model AIC Chi (or F) p-value
fraction of forb seeds (Marone et al., 2017). So, its abundance might
S. multicolor Large + Small + RE 181.7 0.10 0.754 remain unchanged or even increase if there are sufficient alternative
Large + RE 179.8 40.75 < 0.001 seeds in grazed areas (e.g. in this study total seed availability raised
RE 218.6
numerically by 90% (Telteca), 150% (Ñacuñán), and 600% (Alvear) in
M. torquatus Large + Small – F = 0.65 0.424
Large – F = 24.3 < 0.001 the grazed sites for birds able to feed on forb as well as grass seeds in
∼1 – comparison to birds that only feed on grass seeds). Its abundance,
Z. capensis Large + Small 477.6 9.3 0.002 however, may also diminish because a specialist –even an expanding
Large 484.8 20.6 < 0.001 one– will always be more conditioned by its preferred food than a true
∼1 503.4
P. carbonaria Large + Small + RE 206.7 0.04 0.839
generalist species. Thus, inconsistent positive, negative or neutral ef-
Large + RE 204.8 9.1 0.003 fects of cattle grazing on Z. capensis may be expected depending on how
∼RE 211.8 much the bird relaxes its preferences, and on the contingent arrange of
D. diuca Large + Small + RE 301.1 0.06 0.802 available seeds in the field.
Large + RE 299.2 0.82 0.365
Feeding behavior could also account for previous results on win-
∼RE 298.0
tering seed-eating bird responses to domestic grazers in the Reserve of
Ñacuñán. Gonnet (2001) compared bird abundances in the reserve and
the sites (Tables 3 and 4). The abundance of large grass seeds was an overgrazed paddock located in its vicinity in the winters of 1995 and
sufficient to predict the abundances of S. multicolor, M. torquatus and P. 1996, and the fall of 1996. He reported reductions of perennial grasses
carbonaria (Table 3). For the first two species models with different and total grass seeds in the grazed area, where S. multicolor, M. tor-
random structures and error distributions (with better or worse fit) gave quatus, and P. carbonaria diminished on all three occasions, but Z. ca-
similar estimates, whereas for P. carbonaria the results were not as pensis did so on only one occasion. As was also the case in our study,
consistent and the estimation of the best model showed a little more wintering grass seed specialists diminished consistently when facing
uncertainty than for the other species (see confidence intervals in grass seed reductions, whereas Z. capensis showed higher resilience to
Table 4). The best model for predicting the abundance of Z. capensis domestic grazing. If forb seeds were occasionally abundant in the
included the abundance of large grass seeds and small grass/forb seeds grazed area of Gonnet’s (2001) study, this could explain why Z. capensis
(Tables 3 and 4). No model including the abundance of seeds could remained there in two out of three winters.
predict the abundance of D. diuca (Table 3). Although previous results (Cueto et al., 2006; Camín et al., 2015)
pointed that D. diuca is an expanding specialist species which prefer
large grass seeds but can consume small grass and forb seeds when the
4. Discussion
preferred seeds are scarce (Marone et al., 2008, 2017), its abundance
did not track the number of seeds of any functional group in this study.
Overall, cattle grazing did not affect the cover of shrubs and trees in
This lack of response may rise simply because D. diuca behaved in-
the central Monte desert, but diminished grass cover, the number of
consistently in different localities, perhaps following other important
panicles, and of large and small grass seeds and forb seeds in the soil
environmental variables. Certainly, more studies are needed to under-
seed bank. As we expected from bird’s feeding behavior, the decrease of
stand the way cattle grazing affect the abundance of this species, but it
large grass seeds correctly predicted the reduction in the abundance of
is worth noting that predicting numerical responses of species in loca-
the three grass seed specialist passerines (S. multicolor, M. torquatus, and
tions that are at the limit of their breeding-wintering distribution, as is
P. carbonaria). Further, as we also expected from feeding behavior, the
the case for D. diuca (Jaramillo, 2011), may be difficult.
decrease of large grass seeds but also of small grass and/or forb seeds
Given the predictive capacity in 80% of the species assessed, our
accounted for the reductions of Z. capensis. Decreases in seed avail-
results invite to consider the knowledge of seed resource dynamics and
ability due to cattle grazing correctly predicted reductions in the

Table 4
Estimates for the best generalized linear models (GLM) or generalized linear mixed models (GLMM) according to Table 2. Models estimate the effect of different types
of seeds (fixed effects) on the overall counts of five species of birds over three years (n = 8 for each site) in sites under none or low and high grassing pressure in three
localities (random effect) of the Monte Desert, Mendoza, Argentina. Coefficients for the effect of seeds and significant p-values are in bold.
Species Int. Large grass seeds Small grass seeds Locality (SD)

Coef. Lower 95% CL Upper 95% CL Z p-value Coef. Lower 95% CL Upper 95% CL Z p-value

S. multicolor −0.78 1.58 1.07 2.09 6.06 < 0.001 0.60

M. torquatus 1.03 1.69 0.94 2.46 4.38 < 0.001 –
Z. capensis* 2.23 1.52 0.92 2.16 4.74 < 0.001 0.53 0.19 0.89 3.10 0.002 –
P. carbonaria −0.35 1.62 0.59 2.72 3.00 0.003 0.11

* Including forb seeds instead of small grass seeds give the following estimates: Intercept = 1.81; Large grass seeds coefficient (lower 95% CL, upper 95% CL, p-
value) = 1.49 (0.89, 2.14, < 0.001); Forb seeds coefficient (lower 95% CL, upper 95% CL, p-value) = 0.10 (0.04, 0.17, 0.001).

7
M.C. Sagario, et al.
bird foraging behavior to base conservation and management practices
in the Monte. Could these results be just contingent to our data set?
Deepening the review of the specialized literature may help decide
about this crucial question. Bird species that depend heavily on seeds in
their wintering grounds, including those in our study, usually have a
more mixed diet, composed of fruits, arthropods and seeds, during the
breeding season in the South American deserts (Lopez de Casenave
et al., 2008; Milesi et al., 2008; Sánchez and Blendinger, 2014) as well
as in those of North America (Rodewald, 2015). If feeding behavior is a
good predictor of bird responses to grazing, we can predict that birds
will respond less severely to grazing (or will not respond at all) during
the spring and summer because the birds have a wider food resource
base in the breeding season than in the winter. In agreement with this
expectation, Milesi et al. (2002) found a similar abundance of the total
seed-eating bird guild in disturbed and undisturbed areas during two
springs (1988, 1995) and summers (1989, 1996) in the central Monte
desert. Also at the guild level, Blendinger and Ojeda (2001) reported a
positive association between the abundance of perennial grass seeds
and seed-eating birds in winter in localities of the Monte, such as Tel-
teca and El Balde (see Table 2 in Blendinger and Ojeda, 2001), but no
association at all in the spring. Finally, Gonnet’s (2001) species-specific
data for the breeding season showed that S. multicolor did not differ
between conditions or prevailed in the grazed paddock on three out of
four occasions, M. torquatus and D. diuca on two out of four occasions,
and Z. capensis on all four occasions. Only P. carbonaria always pre-
vailed in the reserve in spring or summer (see Fig. 3 in Gonnet, 2001).
Similar seasonal habitat shifts also appear to occur in North
America. According to changes in the density of nine bird species in
desert grasslands and grass steppes (Passerculus sandwichensis,
Aimophila cassini, Ammodramus savannarum, A. bairdii, Calcarius ornatus,
Pooecetes gramineus, Eremophila alpestris, Chondostes grammacus,
Amphispiza bilineata) for which we found information in the literature
on both the wintering and breeding abundance during similar periods,
we classified them as responding negatively, positively or being un-
responsive to grazing in winter sampling (Grzybowski, 1983; Bock
et al., 1984; Bock and Bock, 1999; Gordon, 2000), as well as in spring-
summer sampling (Bock and Webb, 1984; Medin, 1986; Davis et al.,
1999; Powell, 2008; Johnson et al., 2011; Powell and Busby, 2013;
Richardson et al., 2014). Following a “vote-counting” approach
(Gurevitch and Hedges, 1993), we could assess 16 comparisons for
winter sampling, and 21 for the spring-summer. For winter samplings,
we considered that the density of one bird species was higher in one
environmental condition than in the other if the result was statistically
established by the authors in the original paper (Bock et al., 1984;
Gordon, 2000), or the species considered reduced abundance by at least
50% in most years analyzed (Grzybowski, 1983; Bock and Bock, 1999).
For spring-summer samplings, we also employed the results of statistic
hypothesis testing when they were available (Bock and Webb, 1984;
Medin, 1986; Davis et al., 1999; Powell, 2008; Powell and Busby, 2013;
Richardson et al., 2014). For information in Johnson et al. (2011) we
compared the “high grazing intensity” and “control” sites and decided
on the grazing effects based on the consistency of the tendencies during
the four years evaluated. The results confirmed the expectations. In
winter, bird species prevailed in the ungrazed condition in 70% of the
contrasts, and in the grazed condition in only 12% of them. By contrast,
in spring-summer the same bird species predominated in the ungrazed
condition in only 24% of the comparisons, but 47% did so in grazed
areas. The interplay of resource availability and switching diets of seed-
eating birds allows predict a significant fraction of their numerical re-
sponses to grazing in arid and semiarid areas of South as well as North
America.
Coincident patterns of seasonal habitat shift by birds in different
continents, described by different authors for different localities, sup-
port the assumption that resource availability and feeding behavior
could be used to predict bird responses to habitat disturbance. Trophic
ecology was also pertinent to predict reductions in the abundance of
8
Agriculture, Ecosystems and Environment 289 (2020) 106736
seed-eating birds that rely largely on grass seeds feeding in areas where
grasses were progressively excluded by shrubs at Kalahari (Seymour
and Dean, 2010), and the responses of bird species to grazing in grassy
eucalypt woodlands in Australia (Martin and Possingham, 2005).
However, factors other than the availability of seeds may be invoked as
affecting numerical responses of wintering seed-eating birds to habitat
disturbances (Díaz and Tellería, 1994). For example, the presence of
dungs and bare soil in grazed sites may favor carabids and other insects,
which could be eventually incorporated into the diet of seed-eating
birds. Or the abundance of the bird predators (e.g. raptors) may differ
between grazing conditions. None of these options seems likely for our
study because in the same locations and years we did not register
marked differences in abundances between sites for insectivorous birds
(which should be the main beneficiaries of any putative increase in
arthropod prey availability), or birds of prey (Sagario, M.C., Zarco, A.,
Cueto, V.R., Marone, L.; unpublished data). Moreover, even if there are
unmeasured factors that may favor or reduce seed-eating bird abun-
dances at grazed sites, four of the five species showed numerical re-
ductions associated with changes in seed availability owed to grazing
pressure, and these reductions could be predicted and explained based
on their food preferences and diet.
The main effect of grazing on vegetation is often the reduction of the
grass layer (Bertiller, 1992; O’Connor and Pickett, 1992; Bock and
Bock, 1999; Seymour and Dean, 2010; Pol et al., 2014), and grass seed
reserves (Bertiller, 1992; Loydi et al., 2012; Frank et al., 2013; Pol et al.,
2014). Prediction and understanding of the effect of cattle grazing on
seed-eating animals based on natural history may be therefore expected
in semiarid grasslands worldwide, where the level of key seed resources
can be used as “technological knobs” (i.e. factors that can be manipu-
lated with a defined goal and without which eco-technology is not
possible; Marone and Bunge, 1998). Conservative range management
should consider, and even manipulate, the level of the seeds preferred
by wildlife. The use of plants other than grasses with large seeds as
fodder for domesticated livestock or for the restoration of overgrazed
areas, for example, might have negative consequences on most seed-
eating bird populations, and should consequently be avoided. Further,
grazed grasslands should be rested from grazing on a rotational basis so
that grasses, especially those whose seeds are preferred by birds, can
seed. Finally, the focus on natural history traits led to the conciliation of
several apparently contradictory results (e.g. an opposite response of
the same bird species to cattle grazing in winter and summer), identi-
fying the environmental conditions under which ecological results are
reproducible (Marone et al., 2019).
Declaration of Competing Interest
We wish to confirm that there are no known conflicts of interest
associated with this publication and there has been no significant fi-
nancial support for this work that could have influenced its outcome.
Acknowledgements
This paper has benefited from discussion with Javier Lopez de
Casenave, and suggestions made by four anonymous reviewers. Field
work was supported by Agencia Nacional de Promoción Científica y
Tecnológica [grant Pict 2013 2176] and Consejo Nacional de
Investigaciones Científicas y Técnicas [grant PIP 2012 469], both from
Argentina. This is contribution number 100 of the Desert Community
Ecology Research Team (Ecodes) of IADIZA Institute and FCEyN –
Universidad de Buenos Aires.
References
Bates, D., Maechler, M., Bolker, B., Walker, S., 2015. Fitting linear mixed-effects models
using lme4. J. Stat. Softw. 67, 1–48.
Bertiller, M.B., 1992. Seasonal variation in the seed bank of a Patagonian grassland in
M.C. Sagario, et al.
relation to grazing and topography. J. Veg. Sci. 3, 47–54.
Blendinger, P.G., Ojeda, R.A., 2001. Seed supply as a limiting factor for granivorous bird
assemblages in the Monte Desert. Argentina. Aust. Ecol. 26, 413–422.
Bock, C.E., Bock, J.H., 1999. Response of winter birds to drought and short-duration
grazing in Southeastern Arizona. Cons. Biol. 13, 1117–1123.
Bock, C.E., Webb, B., 1984. Birds as grazing indicator species in Southeastern Arizona. J.
Wildl. Manage. 48, 1045–1049.
Bock, C.E., Bock, J.H., Kenney, W.R., Hawthorne, V.M., 1984. Responses of birds, rodents
and vegetation to livestock exclosure in a semidesert grassland site. J. Range.
Manage. 37, 239–242.
Camín, S.R., Cueto, V.R., Lopez de Casenave, J., Marone, L., 2015. Exploring food pre-
ferences and the limits of feeding flexibility of seed-eating desert birds. Emu 115,
261–269.
Camín, S.R., Martín-Albarracín, V., Jefferies, M., Marone, L., 2016. Do neophobia and
dietary wariness explain ecological flexibility? An analysis with two seed-eating birds
of contrasting habits. J. Avian Biol. 47, 245–251.
Cesca, E.M., Villagra, P.E., Alvarez, J.A., 2014. From forest to shrubland: Structural re-
sponses to different fire histories in Prosopis flexuosa woodland from the Central
Monte (Argentina). J. Arid Environ. 110, 1–7.
Cingolani, A.M., Anchorena, J., Stoffella, S., Collantes, M., 2002. A landscape-scale model
for optimal management of sheep grazing in the Magellanic steppe. Appl. Veg. Sci. 5,
159–166.
Cueto, V.R., Marone, L., Lopez de Casenave, J., 2001. Seed preferences by birds: effects of
the design of feeding-preference experiments. J. Avian Biol. 32, 275–278.
Cueto, V.R., Marone, L., Lopez de Casenave, J., 2006. Seed preferences in sparrow species
of the Monte desert: implications for seed-granivore interactions. Auk 123, 358–367.
Cueto, V.R., Milesi, F.A., Sagario, M.C., Lopez de Casenave, J., Marone, L., 2011.
Distribución geográfica y patrones de movimiento de la Monterita canela (Poospiza
ornata) y el Yal Carbonero (Phrygilus carbonarius) en Argentina. Ornitol. Neotrop. 22,
483–494.
Davis, S.K., Duncan, D.C., Skeel, M., 1999. Distribution and habitat associations of three
endemic grassland songbirds in Southern Saskatchewan. Wilson Bull. 111, 389–396.
Díaz, M., Tellería, J.L., 1994. Predicting the effects of agricultural changes in central
Spanish croplands on seed-eating overwintering birds. Agric. Ecosyst. Environ. 49,
289–298.
Díaz, S., Lavorel, S., McIntyre, S., Falczuk, V., Casanoves, F., Milchunas, D.G., 2007. Plant
trait responses to grazing –a global synthesis. Glob. Change Biol. 13, 313–341.
Ehlers-Smith, Y.C., Ehlers-Smith, D.A., Seymour, C.L., Thébault, E., van Veen, F.J.F.,
2015. Response of avian diversity to habitat modification can be predicted from life-
history traits and ecological attributes. Landsc. Ecol. 30, 1225–1239.
Emlen, J.T., 1977. Estimating breeding season bird densities from transect counts. Auk
94, 455–468.
Fontaine, A.L., Kennedy, P.L., Johnson, D., 2004. Effects of distance from cattle water
developments on grassland birds. J. Range Manage. 57, 238–242.
Fox, L.R., Morrow, P.A., 1981. Specialization: species property or local phenomenon?
Science 211, 887–893.
Frank, A.S.K., Dickman, C.R., Wardle, G.M., Greenville, A.C., 2013. Interactions of
grazing history, cattle removal and time since rain drive divergent short-term re-
sponses by desert biota. PLoS One 8, e68466.
Gonnet, J.M., 2001. Influence of cattle grazing on population density and species richness
of granivorous birds (Emberizidae) in the arid plain of the Monte, Argentina. J. Arid
Environ. 48, 569–579.
González Loyarte, M.M., Rodeghiero, A.G., Buk, E., Trione, S., 2000. Análisis comparativo
de dos comunidades en el bosque de Prosopis flexuosaDC. del NE de Mendoza,
Argentina. Multequina (Argentina) 9, 75–89.
Gordon, C.E., 2000. Fire and cattle grazing on wintering sparrows in Arizona grasslands.
J. Range Manage. 53, 384–389.
Gordon, D., 2011. The fusion of behavioral ecology and ecology. Behav. Ecol. 22,
225–230.
Grzybowski, J.A., 1983. Sociality of grassland birds during winter. Behav. Ecol. Sociobiol.
13, 211–219.
Gurevitch, J., Hedges, L.V., 1993. Meta-analysis: combining the results of independent
experiments. In: Scheiner, S.M., Gurevitch, J. (Eds.), Design and Analysis of
Ecological Experiments. Chapman and Hall, New York, pp. 378–398.
Heller, R., 1980. On optimal diet in a patchy environment. Theor. Popul. Biol. 17,
201–214.
del Hoyo, J., Elliott, A., Christie, D.A., 2011. Handbook of the Birds of the World Vol. 16
Lynx Edicions, Barcelona.
Isacch, J.P., Cardoni, D.A., 2011. Different grazing strategies are necessary to conserve
endangered grassland birds in short and tall salty grasslands of the flooding pampas.
Condor 113, 724–734.
Jaramillo, A., 2011. Common Diuca-Finch Diuca Diuca. In: del Hoyo, J., Elliott, A.,
Christie, D.A. (Eds.), Handbook of the Birds of the World, vol. 16. Lynx Edicions,
Barcelona.
Agriculture, Ecosystems and Environment 289 (2020) 106736
Johnson, T.N., Kennedy, P.L., DelCurto, T.V., Taylor, R.V., 2011. Bird community re-
sponses to cattle stocking rates in a Pacific Northwest bunchgrass prairie. Agric.
Ecosyst. Environ. 114, 338–346.
Kutt, A.S., Martin, T.G., 2010. Bird foraging height predicts bird species response to
woody vegetation change. Biodivers. Conserv. 19, 2247–2262.
Lopez de Casenave, J., 2001. Estructura gremial y organización de un ensamble de aves
del desierto del Monte, [Dissertation, Universidad de Buenos Aires]. (Accessed 10
February 2019). http://digital.bl.fcen.uba.ar/Download/Tesis/Tesis_3319_
LopezdeCasenave.pdf/.
Lopez de Casenave, J., Cueto, V.R., Marone, L., 2008. Seasonal dynamics of guild struc-
ture in a bird assemblage of the central Monte desert. Basic Appl. Ecol. 9, 78–90.
Loydi, A., Zalba, S.M., Distel, R.A., 2012. Viable seed banks under grazing and exclosure
conditions in montane mesic grasslands of Argentina. Acta Oecol. 43, 8–15.
Marone, L., Bunge, M., 1998. La explicación en ecología. Boletín de la Asociación
Argentina de Ecología 7, 35–37.
Marone, L., Camín, S.R., Cueto, V.R., 2015. Context dependent foraging by seed-eating
birds does not necessarily mean low ecological predictability. Can. J. Zool. 93,
353–359.
Marone, L., Lopez de Casenave, J., Milesi, F.A., Cueto, V.R., 2008. Can seed-eating birds
exert top-down effects on grasses of the Monte desert? Oikos 117, 611–619.
Marone, L., Olmedo, M., Valdés, D.Y., Zarco, A., Lopez de Casenave, J., Pol, R.G., 2017.
Diet switching of seed-eating birds wintering in grazed habitats of the central Monte
Desert, Argentina. Condor 119, 673–682.
Marone, L., Lopez de Casenave, J., González del Solar, R., 2019. The synthetic thesis of
truth helps mitigate the "reproducibility crisis" and is an inspiration for predictive
ecology. Revista de Humanidades de Valparaíso 14 In press.
Martin, T.G., Possingham, H.P., 2005. Predicting the impact of grazing on birds using
foraging height data. J. Appl. Ecol. 42, 400–408.
Medin, D.E., 1986. Grazing and passerine breeding birds in a great basin low-shrub de-
sert. Great Basin Nat. 46, 567–572.
Milesi, F.A., Lopez de Casenave, J., Cueto, V.R., 2008. Selection of foraging sites by desert
granivorous birds: vegetation structure, seed availability, species-specific foraging
tactics, and spatial scale. Auk 125, 473–484.
Milesi, F.A., Marone, L., Lopez de Casenave, J., Cueto, V.R., Mezquida, E.T., 2002.
Gremios de manejo como indicadores de las condiciones del ambiente: un estudio de
caso con aves y perturbaciones del hábitat en el Monte central, Argentina. Ecol. Aust.
12, 149–161.
O’Connor, T.G., Pickett, G.A., 1992. The influence of grazing on seed production and seed
banks of some African savanna grasslands. J. Appl. Ecol. 29, 247–260.
Pol, R.G., Sagario, M.C., Marone, L., 2014. Grazing impact on desert plants and soil seed
banks: implications for seed-eating animals. Acta Oecol. 55, 58–65.
Powell, A.F.L.A., 2008. Responses of breeding birds in tallgrass prairie to fire and cattle
grazing. J. Field Ornithol. 79, 41–52.
Powell, A.F.L.A., Busby, W.H., 2013. Effect of grassland management on breeding birds at
the Western edge of the tallgrass prairie ecosystem in Kansas. Nat. Areas J. 33,
130–138.
R Core Team, 2016. R: A Language and Environment for Statistical Computing. R
Foundation for Statistical Computing, Vienna, Austria (Accessed 10 February 2019).
https://www.R-project.org/.
Richardson, A.N., Koper, N., White, K.A., 2014. Interactions between ecological dis-
turbances: burning and grazing and their effects on songbird communities in northern
mixed-grass prairies. Avian Conserv. Ecol. 9, 5.
Ríos, J.M., Mangione, A.M., Marone, L., 2012. Effects of nutritional and anti-nutritional
properties of seeds on the feeding ecology of seed-eating birds of the Monte Desert.
Argentina. Condor 114, 44–55.
Rodewald, P. (Ed.), 2015. The Birds of North America. Cornell Laboratory of Ornithology,
Ithaca (Accessed 10 February 2019). http://bna.birds.cornell.edu/BNA/.
Sagario, M.C., Cueto, V.R., Lopez de Casenave, J., 2014. Movement patterns of three
species of sparrow in the central Monte desert: differences between and within spe-
cies. Emu 114, 268–276.
Sánchez, R., Blendinger, P.G., 2014. Trophic ecology of the Ringed Warbling-Finch
(Poospiza torquata) in Neotropical semi-arid scrublands. Emu 114, 229–233.
Seymour, C.L., Dean, W.R.J., 2010. The influence of change in habitat structure on the
species composition of bird assemblages in the southern Kalahari. Aust. Ecol. 35,
581–592.
Verner, J., 1985. Assessment of counting techniques. Curr. Ornithol. 2, 247–302.
von Müller, A.R., Cingolani, A.M., Vaieretti, M.V., Renison, D., 2012. Estimación de carga
bovina localizada a partir de frecuencia de deposiciones en un pastizal de montaña.
Ecol. Aust. 22, 178–187.
Venables, W.N., Ripley, B.D., 2002. Modern Applied Statistics with S, fourth ed. Springer,
New York.
Zuur, A.F., Ieno, E.N., Walker, N.J., Saveliev, A.A., Smith, G.M., 2009. Mixed Effects
Models and Extensions in Ecology with R. Springer Science + Business Media, New
York.