ROLE OF LANDSCAPE ELEMENTS ON RECENT DISTRIBUTIONAL

EXPANSION OF EUROPEAN STARLINGS (STURNUS VULGARIS)
IN AGROECOSYSTEMS OF THE PAMPAS, ARGENTINA

EMMANUEL ZUFIAURRE,1 AGUSTIN ABBA,2

DAVID BILENCA,1 AND MARIANO CODESIDO1,3

ABSTRACT.—Previous studies of European Starlings in Argentina have focused on identifying biological aspects

correlated with establishment of new populations in urban and suburban areas. Starlings have recently invaded rural areas

in the Pampas. To understand the factors involved in the recent expansion of European Starlings into these rural habitats,

we investigated how expansion patterns were associated with season, proportion of crop fields, distance to woodlots, and

distance to small and big urban centers in agroecoystems of the Pampas. We surveyed 392 fields during 2011–2013 to

collect data on presence of starlings and landscape features. We found that the range of European Starlings has expanded by

a total area of ,65,000 km2 since 2005, at a linear range expansion rate of 22.2 km per year. Generalized linear mixed

model analysis revealed that presence of European Starlings was significantly related with reduced distances to nearest

small urban area. Our findings indicate that range expansion of European Starlings into rural areas of Argentina may follow

a neighborhood diffusion pattern, by which well-established populations act as sources of individuals that disperse short

distances into nearby favorable areas. In absence of human control, this species is expected to continue its spread and

population increase. Received 22 April 2015. Accepted 22 November 2015.

Key words: agriculture, biogeography, biological invasions, Neotropics, Sturnidae.

Landscapes in South America and elsewhere

have been heavily modified for agricultural pro-

duction (Bruggers et al. 1998, Bomford and

Sinclair 2002, Codesido et al. 2015). The Pampas

of central Argentina is one of the largest agricul-

tural regions of the world (Soriano et al. 1992). The

pre-European Pampas was homogeneous grass-

land; however, native vegetation has been in-

creasingly replaced by crops through the twentieth

century (Baldi and Paruelo 2008). Even though

originally the Pampas had no treed vegetation,

woodlands with both native and exotic tree species

have become established along riparian habitat and

roadsides and were intentionally planted around

railway stations, near rural buildings, on ranches as

shading woodlots for cattle, and in urban areas

(Ghersa et al. 2002, Zalba and Villamil 2002).

The introduction of trees to the Pampas was

followed by the expansion of opportunistic birds

that nest on them, such as doves, pigeons, and

parakeets (Daguerre 1936, Murton et al. 1974,

Bucher and Aramburú 2014, Codesido et al.
2015). These birds are now among the most
abundant species in the bird assemblage of the
Pampas and are widely regarded as avian crop
pests (Bruggers et al. 1998, Codesido et al. 2012).
Pest birds rely heavily on herbaceous plants,
bushes, and trees for feeding, nesting, and shelter
(Warburton and Perrin 2006, MacLeod et al.
2011, Bucher and Aramburú 2014). Most previous
studies have focused on how increases in avian
crop pest populations are associated with land-
scape characteristics, such as crop types and the
presence of rangelands, woodlots, urbanization,
roads and railway networks, since these modi-
fied environments provide important resources for
these pest species (Bucher and Aramburú 2014,
Codesido et al. 2015). Less research effort has
focused on how avian crop pests are expanding
their ranges or how non-native birds are invading
these agricultural landscapes and presenting new
agricultural threats (Bruggers et al. 1998, Bucher
and Aramburú 2014).

One of the most widespread invasive birds
throughout the world is the European Starling
(Sturnus vulgaris), which has many traits associ-
ated with introduction success (Duncan et al. 2003)
and has been listed as one of the world’s 100 worst
invasive species (Lowe et al. 2004). Originally
native to Europe and Asia, it has expanded its range
over the past hundred years and has established
viable populations on Australia (introduced in

1 Grupo de Estudios sobre Biodiversidad en Agroecosiste-

mas (GEBA), Departamento de Biodiversidad y Biologı́a

Experimental, Facultad de Ciencias Exactas y Naturales,

Universidad de Buenos Aires and IEGEBA (UBA-

CONICET), Ciudad Universitaria, Pabellón II, 4u Pisa,

(C1428EHA), Ciudad Autónoma de Buenos Aires, Argentina.
2 Centro de Estudios Parasitológicos y de Vectores

(CEPAVE) CCT - CONICET - LA PLATA- UNLP. La

Plata (B1902CHX), Buenos Aires, Argentina.
3 Corresponding author; e-mail: mcodesido@ege.fcen.uba.ar

The Wilson Journal of Ornithology 128(2):306–313, 2016

306



1856), New Zealand (1862), North America
(1890), South Africa (1899), and Pacific and
Caribbean Islands (Flux and Flux 1981, Feare
1984). More recently, European Starlings were
accidentally released in Argentina, and were first
recorded in wooded urban areas of Buenos Aires
city in 1987 (Pérez 1988). As of 2005, the range
of European Starlings bordered the Pampas and
ran along the Atlantic coast up to 30 km inland
(Peris et al. 2005). Although its early dispersal
history was strongly associated with urban areas
(Peris et al. 2005, Rebolo Ifran and Fiorini 2010,
Girini et al. 2014), starlings have recently begun
spreading into suburban and rural environments
(Jensen 2008), and have recently been recorded
in rural Pampas (EZ, pers. obs.).

The expansion of starlings into rural areas is
of particular concern, because starlings are
avian crop pests and large populations can cause
significant economic losses in agriculture
(Campbell et al. 2015). Starlings eat a wide range
of seeds, grains, and fruits from both native and
cultivated species (Feare et al. 1992, Bomford and
Sinclair 2002, Pimentel et al. 2005), and also feed
on a variety of invertebrates such as leatherjackets
(Diptera), earthworms (Lumbricidae), caterpillars
(Orthoptera), pillbugs (Isopoda), and Lepidoptera
(Westerterp et al. 1982, Moore 1986). Starlings
use holes for nesting but are not primary excavators
(Feare 1984). In Argentina, starlings use a wide
array of facilities as nest sites, although most nests
are in holes excavated by woodpeckers (Colaptes
spp.), mostly in exotic species of old trees, such
as elms (Ulmus spp.), poplars (Populus spp.),
eucalyptus (Eucalyptus spp.), and oaks (Quercus
spp.), which tend to be planted in both urban and
rural areas (Peris et al. 2005). Given the invasive
characteristics of European Starlings, it is likely
that they will become abundant in the future and
may significantly impact agricultural production
and native ecosystems. Characterizing the pattern
of spread of invasive species constitutes a neces-
sary step in managing biological invasions (Hulme
2006).

The goal of this work was to document the recent
invasion of European Starlings into agroecosys-
tems in the Argentinean Pampas and understand
the landscape factors involved with range expan-
sion into more rural areas. Specifically, our aims
were to (1) estimate the size and rate of rural range
expansion of European Starlings in the Pampas
agroecosystems and (2) examine associations
between presence of European Starlings and

landscape features such as land-use, distance
to woodlots, and distance to small and big urban
centers.

MATERIAL AND METHODS

Study Area.—Our study area in the Pampas
region of central Argentina extends over about
225,000 km2 (500 km from north to south, 450 km
from east to west; 33–39uS, 57–63u W; Fig. 1).
The region is almost flat, with only a few hills
and rocky outcrops at isolated sites. Mean annual

temperature is ,15uC, with warm summers and
cool winters (Jan mean temperature ranges 21.5–
23.5uC, Jul mean temperature ranges 7.5–9.5uC);
mean annual precipitation ranges 800–1000 mm
(Soriano et al. 1992).

The natural vegetation is a gramineous steppe
dominated by grasses such as Nasella, Piptochae-
tium, Aristida, Melica, Briza, Bromus, Eragrostis
and Poa, intermingled with prairies, marshes and
other edaphic communities (Soriano et al. 1992).
More recently, the Pampas region has been used for

intensive crop production not managed by tilling or
cattle grazing (Bilenca et al. 2012). Planted stands
of tall trees are new to the Pampas and have added
structural complexity to the grasslands (Zalba and
Villamil 2002).

Surveys of European Starlings.—From Decem-
ber 2011 to June 2014, we recorded European
Starlings through both systematic and opportunis-
tic surveys. We conducted systematic surveys at
25 sites distributed evenly throughout our study

area (Fig. 1). Sites were at least 40 km apart and
had varying proportions of land under crop
production and livestock use. In each of these
sites, we selected four independent fields, two crop
stubbles, and two livestock paddocks. We did four
surveys in each site: two during spring-summer

(2011–12 and 2012–13) and two during autumn
(2012 and 2013). Thus, each site was surveyed
twice each season throughout two years, but each
sampling was carried out in different fields, so
that, we avoided dependence among data. We
surveyed pastures and natural or semi-natural

grasslands in livestock paddocks, and surveyed
winter crop stubble (wheat Triticum aestivun,
barley Hordeum vulgare, rye Secale cereale)
during spring–summer and summer crop stubble
(soybean Glycine max, corn Zea mays, sunflower
Helianthus annuus, and sorghum Sorghum sp.)

during autumn. In total, we surveyed 392 fields
(196 of each land use type; at least 1000 m apart),

Zufiaurre et al. N EUROPEAN STARLING IN PAMPAS AGROECOSYSTEMS 307



since during December 2011 we could not survey
in two sampling sites because of logistical
problems. Bird surveys were carried out 4 hrs after
dawn. Within each field and avoiding boundaries,
we randomly established a transect 700 m long by
100 m wide (Bibby et al. 2000), and walked the
transect for 15 mins while recording the numbers of
European Starlings by transect. We also conducted
opportunistic surveys during extensive traveling by
the authors covering the study area, where we
recorded presence of starlings while driving slowly
along primary and secondary roads.

Surveys of Landscape Attributes on Systematic
Samplings.—For each transect at each field we
recorded: a) the proportion of crop fields surround-
ing the transect (including the field where the
transect was set and the adjacent fields to that
field), b) distance in meters from the center of the
transect to the nearest woodlot (patches with
presence of exotic trees .0.2 ha), c) distance from
the center of the transect to the nearest big urban
area (i.e., with more than 10,000 people), and d)
distance from the center of the transect to the
nearest small urban area (i.e., with less than 10,000

people, train stations, or farmhouses). Distance
variables were measured using Google EarthH
images, expressed in meters.

Statistical Analyses.—To estimate the range
expansion of European Starlings, we compared
our bird distribution data (systematic and opportu-
nistic records) with baseline data reported by
Peris et al. (2005) who compiled data from 1987
to 2004 on the basis of personal observations,
published sources, and field reports provided by
experienced birdwatchers. The linear range expan-
sion for European Starlings was estimated as the
average linear distance between the initial and
final distribution borders between both databases.

To examine the associations between landscape
attributes and presence of European Starlings
at its recent expansion area (northeast and east of
province, this study; n 5 10 sites; 160 transects),
we compared certain landscape attributes at the
27 transects where European Starlings were
recorded with another 27 transects without the
species which we randomly choose from the total
of 133 sampled transects where the species was
absent. We used generalized lineal mixed models

FIG. 1. Study area and locations of sampling sites in agroecosystems of the Pampas, Argentina. Detail of distribution of

European Starlings in Argentina by Peris et al. (2005).

308 THE WILSON JOURNAL OF ORNITHOLOGY N Vol. 128, No. 2, June 2016



(Zuur et al. 2009) in order to analyze the
association between farmland landscape attributes
and presence of European Starlings by field (the
response variable). The explanatory variables were
as follows: season (with two levels, spring-summer
and autumn) was included as categorical variable,
and proportion of crop fields surrounding the
transect, distance to woodlot, distance to large
urban area, and distance to small urban area as
continuous variables. All these variables were
specified as fixed effects, whereas site was treated
as a random effect. The presence of European
Starlings fitted binomial data distribution (pres-
ence/absence) and ‘logit’ link function (Crawley
2007). We employed a backward selection pro-
cedure removing non-significant terms from the
model one by one, in decreasing order of pro-
bability. The stepwise procedure from the saturat-
ed model (i.e., with all explanatory variables)
through a series of simplifications to the mini-
mal adequate model was made on the basis of
deletion chi-squared tests that assessed the
significance of the increase in deviance that
results when a given term is removed from the
current model (Crawley 2007). We checked
the model fit by means of graphical validation
tools for the binomial data distribution (Zuur
et al. 2009). Statistical analyses were carried
out using R software, version 3.1.0 (R Core Team
2014).

RESULTS

Range Expansion.—From December 2011 to
June 2014, starlings were present in 84 surveys
(27 systematic and 57 opportunistic surveys)
throughout the agroecosystems of the Pampas.
We recorded a total of 1141 individuals (composed
of individual records and flocks of up to 110
individuals), 56% in spring-summer (mean 5 11.9
individuals, ES 5 2.2, n 5 54) and 44% in
autumn (mean 5 16.6 individuals, ES 5 5.1,
n 5 30). The estimated average linear distance
between the initial and current distribution for
European Starlings was 221.8 km, with an average
progression from 2005–2014 of 22.2 km per year
(Fig. 2).

Associations between Landscape Elements and
Presence of European Starlings.—Our generalized
linear mixed model showed that presence of
European Starlings was attributed to the distance
to the nearest small urban area (mean 5 6,627 m,
ES 5864 m, n 5 54; Table 1 and 2). The presence
of the species was not significantly associated with

season, or with distance to nearest big urban area
(mean 5 14,329 m, ES 51,534 m, n 5 54), or
with distance to nearest woodlot (mean 5 332 m,
ES 5 30 m, n 5 54), or with crop landscape
(mean 5 53%, SD 5 32%, n 5 54). Closeness
to small urban areas increased the likelihood that
starlings were present. The average distance to the
nearest small urban area in fields with presence of
European Starlings (mean 5 4,359 m, ES 5 90 m,
n 5 27) was almost 51% less than the average
distance in fields without European Starlings
(mean 5 8,895 m, ES 5 300 m, n 5 27).

DISCUSSION

The spread of European Starlings seems to be
still far from finished 28 years after its introduc-
tion in Argentina. Instead, the range expansion is
increasing both in size and rate. Expansion of
European Starlings into the Pampas covers an area
close to 65,000 km2,more than the area already
reported by Peris et al. (2005), with an average
range expansion of 22.2 km per year. This
expansion rate is 296% greater than the average
expansion previously quantified (Peris et al. 2005;
7.5 km per year), although slow if compared
with the 43 km per year of European Starlings in
North America over the twentieth century (Wing
1943). Importantly, this rate might underestimate
the real expansion rate, since European Starlings
were not detected in intensive surveys made during
2006–2008 (Codesido et al. 2012, 2013; Fig. 2),
thus the recent expansion of the species within the
Pampas agroecosystems of Argentina most likely
occurred very recently (i.e., since 2009). Our
results are in agreement with recent records of
European Starlings that have also been reported
opportunistically by several birders in the Pampas
since 2010 (eBird 2015).

Our study revealed that some landscape ele-
ments influenced the process of range expansion by
starlings in the Pampean agroecosystems of
Argentina. European Starlings colonized sites that
were close to small urban areas (small towns, train
stations, or farmhouses). Most of these human
settlements had treed vegetation around houses,
providing suitable nesting sites and roosting, as
well as peridomestic fruit orchards (mulberries,
figs, date palms, etc.) which provide feeding sites.
In addition, urban areas usually act as centers of
well-established populations (Kessel 1953), so
that dispersal of starlings may follow a neighbor-
hood diffusion pattern, by which well-established
populations act as sources of individuals that

Zufiaurre et al. N EUROPEAN STARLING IN PAMPAS AGROECOSYSTEMS 309



travel short distances into nearby favorable areas
(Bucher and Aramburú 2014). This pattern is
consistent with the pattern of diffusion from the
east to the west reported for USA (Wing 1943,

Kessel 1953). Patterns of invasion and range

expansion in the USA have been attributed to the

movement of first-year and non-breeding second-

year starlings during migration or more general

wandering dispersal patterns (Kessel 1953). These

irregular movements, and the apparent random

selection of breeding areas by younger birds, would

be sufficient to account not only for extensions of

the breeding range but also for many of the winter

extensions.

Besides the proximity to small urban areas, we

found no evidence that other landscape elements

may have influenced the expansion of European

TABLE 1. Generalized linear mixed model of the proba-

bility of presence of European Starlings in agroecosystems of

the Pampas, Argentina. N 5 54 transects (27 presences and

27 absences).

Explanatory variables Coefficients 6 E.S. Z P

Intercept 1.211 6 0.6012 2.01 0.044

Distance to nearest

small urban area 20.0002 6 0.00009 2.28 0.029

FIG. 2. Expansion of European Starlings (detail of systematic and opportunistic records) in agroecosystems of the

Pampas, Argentina.

310 THE WILSON JOURNAL OF ORNITHOLOGY N Vol. 128, No. 2, June 2016



Starlings in the rural Pampas region. Climatic

factors may be excluded (at least in terms of the

starlings’ physiological tolerance), because the

climate of the Pampas is within the same boundary

value ranges of the surrounding areas, where

European Starlings were introduced originally.

Even though initial expansion of European

Starlings occurred slowly, the rate of expansion

is currently accelerating, which could indicate that

European Starlings have overcome the lag phase

of the invasion process. Population lag phases

have been defined as a period of slow population

growth followed by a marked increase in the

rate of growth and are common in the establish-

ment and spread of many invasive species (Crooks

2005, Aikio et al. 2010). The occurrence of lag

phases within exotic bird populations makes it

possible for currently rare exotic species to later

explode in numbers and geographic extent

(Aagaard and Lockwood 2014). The current range

expansion pattern indicates that European Star-

lings have left the lag phase of their invasion in

Argentina and may be entering a period of

explosive growth.

Management Implications.—Once introduced

species are established, their control or eradication

is difficult and likely to be highly costly (Kolar and

Lodge 2001, Campbell et al. 2015). European

Starlings are already successfully established in the

Pampas. In the absence of human control, this

species is expected to continue its spread and

population increase, because of its adaptability to

urban and suburban habitats (Lovell 1941, Feare

1984), its nearly omnivorous diet (Moore 1986, Feare

et al. 1992), and its large reproductive output (Feare

1984). This potential for further expansion, together

with its agricultural pest status in its exotic range

(Bomford and Sinclair 2002, Lowe et al. 2004),

should be taken into account when considering the
potential damage it may cause particularly in western
areas of Argentina like the provinces of Mendoza,
San Juan, La Rioja, Catamarca and Salta, where
vineyards and olive groves (which have been heavily
damaged by starlings elsewhere; Bomford and
Sinclair 2002) are among the main agricultural
productions. Moreover, although in Argentina Euro-
pean Starlings build nests in areas environmentally
disturbed (Peris et al. 2005), they may also
represent a novel competitor with native species for
nest sites, although the impact on native biota is much
more difficult to predict (Ingold 1998, Massaro et al.
2013). Considering all the above and the fact that the
earlier intervention is carried out, the better the
economic return on control investment (Campbell et
al. 2015), there is an urgent need to start a control
program of starlings efficiently and effectively in the
Pampas.

ACKNOWLEDGMENTS

Special thanks to the workers and owners of all the

agricultural establishments that allowed access to work on

their property (particularly to Alan Goodall, INTA Perga-

mino, CEPT Nu5 Miranda, Bomberos de Udaquiola, Tatay,

El Haras, Hermanos Laplace, Hinojales, Pelerı́, La Torcacita,

Don Remigio, Santa Elena de Inchauspe, Monte Unión, La

Providencia, Manantiales). We appreciate the review and the

improvements in English usage made by Elizabeth Hobson

through the Association of Field Ornithologists’ program

of editorial assistance. Financial support was provided by

Consejo Nacional de Investigaciones Cientı́ficas y Técnicas

(CONICET PIP 2010-2012 GI 11220090100231), Agencia

Nacional de Promoción Cientı́fica y Tecnológica (BID

PICT2010-1412), Universidad Nacional de La Plata (PPID/

N004), and Universidad de Buenos Aires (UBACyT GC

20020090100070, GC 20020120100018).

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