Glorious past, uncertain present, bad future? Assessing effects of land-use changes on habitat suitability for a threatened farmland bird species
Introduction
Changes in land-use have a global effect (Leemans and Zuidema, 1995, Hurtt et al., 2006) and, interacting with other components of global change, have a strong impact on the structure of biological communities (Sala et al., 2000, Thomas et al., 2004, Rosenzweig et al., 2008, Gil-Tena et al., 2009). Mediterranean ecosystems appear to be especially susceptible to the impacts of global change (Lavorel et al., 1998), because of the high exposure to human activities (Blondel, 2006), the sensitivity to climatic conditions (Peñuelas et al., 2002) and the simultaneous impact of several factors (Sala et al., 2000). The effects on biodiversity of such impacts are largely unknown (Sala et al., 2000), may be non-linear and may display opposite directions (Gil-Tena et al., 2009).
Land abandonment has been an important land-use change in recent decades, being a major consequence of socio-economic changes in highly industrialized countries (Ostermann, 1998). The decrease in farming primarily affects the least productive agricultural land and triggers a recovery of semi-natural vegetation (Sirami et al., 2008). In most of the Mediterranean region, land abandonment has occurred during the last century. This has led to the naturalization and vegetation closure of many areas, thus favouring the spread of forests (e.g. Debussche et al., 1999, Poyatos et al., 2003, Roura-Pascual et al., 2005). The Northern Mediterranean basin has a long-term and complex history of human exploitation, which has largely shaped its landscapes and associated fauna (Blondel, 2006). It has recently experienced fast and widespread agricultural abandonment (Mazzoleni et al., 2004), which appears to be an important cause of species declines, especially in bird communities (Farina, 1995, Farina, 1997). The recovery of semi-natural vegetation that results from agricultural abandonment causes a decrease of open grassland-like habitats and an increase in shrubland and, especially on the long-term, woodland cover (Debussche et al., 1999, Romero-Calcerrada and Perry, 2004). This causes a decline of species tied to open habitats (Preiss et al., 1997, Suárez-Seoane et al., 2002, Sirami et al., 2007), with migrant species associated with open farmland habitats being particularly affected (Sirami et al., 2008). In general, many species of conservation concern in Europe are associated with traditional farm landscapes and the semi-natural habitats they produce and maintain (see e.g. Tucker and Evans, 1997). Such types of landscapes have been increasingly studied in order to understand mechanisms underlying the decline of farmland birds (Benton et al., 2003) and the main steps to be taken for their conservation (Donald et al., 2002).
On the basis of the above, assessments are urgently required on the impact (past, current, and likely future) of environmental change, on animal communities and especially on threatened bird species in the Mediterranean region. Assessments should especially include land abandonment and the consequential fast recovery of forest, further boosted by climate change and frequency of fires (Grigulis et al., 2005).
However, as it is rare to have enough information on past land-uses: (i) it is difficult to assess previous ecological gain or loss, and (ii) future predictions are often built up without a historical base. Although notable exceptions exist (see e.g. Domenech et al., 2005, Ficetola et al., 2009, Fouquet et al., 2010), none of them consider farmland birds, one of the main concerns for conservationists at the beginning of the third millennium.
Among depleted and declining farmland birds, the red-backed shrike Lanius collurio is a typical inhabitant of semi-open habitats created and maintained by ‘traditional’, low-intensity agricultural practices (Fornasari et al., 1997). A recent study revealed that shrike conservation in southern European farmed landscapes depends on a trade-off between ‘intense’ agricultural land use and ‘partial’ land abandonment, which provide complementary resources (Brambilla et al., 2007a). Other research in different European countries has highlighted a similar pattern: red-backed shrikes occupy mostly farmed landscapes characterised by a matrix of fields and grasslands, within which marginal features are interspersed, such as shrubs, hedgerows and small trees (Kuzniak and Tryjanowski, 2000, Vanhinsbergh and Evans, 2002, Karlsson, 2004, Tryjanowski et al., 2007, Golawski and Golawska, 2008, Golawski and Meissner, 2008, Brambilla et al., 2009a). These widely converging findings, coming from different geographical contexts, emphasise how red-backed shrikes are principally tied to an ‘intermediate’ ecological disturbance created by agro-pastoral activities over biological communities and environment. Red-backed shrikes need open areas with low grass provided by pastures, mown grasslands or grassy arable-fields; they also need shrubs or small trees for perching and nesting sites, which are found in areas not directly used for agricultural purposes, or by portions of pasture with a lower grazing pressure. Finally, untilled herbaceous plants favour shrike occurrence by positively affecting the abundance and availability of invertebrate prey; untilled grasslands are often ensured by fallow land or set-aside. When a similar mosaic pattern (which in most European countries does not occur as a ‘natural’ feature but only as a by-product of traditional agricultural land use) is created ‘ad hoc’ by habitat management, shrikes respond positively to this improved habitat quality with a steady population increase, stressing the link between this species and the habitat mosaic created by low-intensity agriculture (Casale et al., 2007, and references therein).
Red-backed shrikes are quite easy to note and locate, thanks to their conspicuous behaviour, and previous spatially explicit models of habitat suitability for this species, based on land use features, lead to valid predictions of species abundance (Brambilla et al., 2009a). The red-backed shrike in the northern Mediterranean basin thus appears to be an ideal model for investigating the effect of habitat changes, caused by land abandonment, on threatened farmland birds.
With this work we model habitat suitability for the study species according to dynamic land-use contexts. We analyse the species’ habitat requirements in a landscape widely shaped by traditional agricultural use, where historical changes in land use and associated vegetation structure are known; then, we explore variations in habitat suitability from the 1950s to the present and predict future habitat suitability in a land-abandonment scenario.
The simultaneous occurrence of a range of neighbouring land-types allows us to investigate how strictly red-backed shrikes are dependent on different types of land use. Such types include land for non-intensive crop production and livestock grazing as well as arable-fields, pastures and woodland that have been abandoned recently or in the past. We formulate a spatially explicit model of habitat suitability for the species according to the current pattern of land-use types. Then we evaluate past habitat suitability, by applying our model to three known past scenarios (dating back to the 1950s), and simulate the development of vegetation communities after complete land abandonment, i.e. the cessation of all agricultural and pastoral activities. We simulate the progressive substitution of fields and pastures by shrubland and tree communities, setting the time and order of habitat succession on the basis of the historical evidence of land-use changes. By estimating past habitat suitability and by simulating the future evolution of land-use cover in an abandonment scenario (quite likely to occur in future decades), we further test the effect of land abandonment on shrike occurrence. Land abandonment has been the major force driving landscape changes in recent years in the study area (Bogliani et al., 2003; see also Fig. 1) and in other mountainous regions.
The analysis of possible future scenarios, calibrated using historical data, allowed us to predict the effects of land abandonment on this declining species. Red-backed shrike could also be used as a model for entire biological communities of low-intensity agricultural areas, at least in northern Italy, where it is an indicator of high biodiversity (Brambilla et al., 2009a, Casale and Brambilla, 2009). Since land-use changes (like the ones we consider here), represent a common phenomenon in most of southern European and Mediterranean mountainous areas, we believe that our study species/case may have a more general interest, and may often be applicable to the species-rich (and conservation priority) biological communities of traditionally farmed landscapes.
Section snippets
Study area
The study area (18,040 ha) is located in the northern Apennines (Italy, Lombardy, province of Pavia, Fig. 1), roughly on the boundary of the Mediterranean and Eurosiberian regions. This area holds a large diversity of farmland birds (Bogliani et al., 2003) and important populations of many conservation-concern species (e.g. Negri et al., 2005, Brambilla et al., 2008, Brambilla et al., 2009b). The red-backed shrike is the only shrike species regularly breeding in the area; therefore, the
Habitat association
The AIC-based model selection and the two stepwise backward procedures provided nearly identical results, with all three methods leading to the same model. The only exception was for permanent grassland, which resulted marginally significant in stepwise procedures (P ∼ 0.1 under both approaches) and which was therefore left out from those models at the last step. However, the model including permanent grasslands seemed to be more meaningful, as many of the cultivated fields within shrikes’
Moderate farming creates key habitat resources destroyed by land abandonment
Red-backed shrike territories include a compact mosaic of different habitats (Brambilla et al., 2009a). This is clearly pointed out by the habitat preference model, which indicates the need for both grassland-like habitats and for shrub cover. Fields, permanent grasslands and shrub habitats all have positive effects on the probability of shrike occurrence, while, on the other hand, the only variable negatively affecting shrike occurrence is broadleaved woodland, by far the commonest forest type
Acknowledgments
We are very grateful to G. Bottari, P. Colombani, G. Ferrario, P. Lenna, A. Martinoli, D. Moreno, F. Piccarolo, F. Picco, S. Ronchi, P. Siccardi, G. Tosi, P. Volonterio for helpful comments and cooperation. Four anonymous reviewers and the editor B. Söderström provided helpful suggestions on a first draft of the ms. N. Sills kindly revised the English text. This study has been funded by FLA and by DGQA of Lombardy Region, and is part of FC’s PhD.
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