NICOLS SEOANE

Habitat use of semi-feral cattle

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First records on habitat use of semi-feral cattle in southern forests: topography reveals

more than vegetation

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Nicols Seoane

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Laboratorio Ecotono, Universidad Nacional del Comahue, Quintral 1250, (8400) Bariloche,

Argentina. (Email: nicosaon80@gmail.com) Tel. +54 (02944) 423374

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ABSTRACT

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While the presence of cattle in forests is quite common, how they use the habitat is often

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overlooked. When this is examined, most studies focus on measurements of the vegetation

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variables influencing habitat selection. This current study provides a suitable model to study

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habitat use by livestock in forested areas. The model was applied to data from semi-feral cattle in

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order to obtain the first description of their habitat use in southern forests. Furthermore, the

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model accounted for individual variability, and hinted at population patterns of habitat use. The

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positions of four individual animals with GPS collars were recorded during four months in a

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Nothofagus (southern beech) forest in Patagonia (Argentina). By projecting these GPS location

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data into a geographical information system (GIS), a resource selection probability function

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(RSPF) that considers topographic and vegetation variables was built. The habitat use of semi-

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feral cattle in southern beech forests showed a large interindividual variability, but also some

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similar characteristics which enable a proper description of patterns of habitat use. It was found

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that habitat selection by cattle was mainly affected by topographic variables such as altitude and

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the combination of slope and aspect. In both cases the variables were selected below average

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relative to availability, suggesting a preferred habitat range. Livestock also tended to avoid areas

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of closed shrublands and showed a slight preference for pastures (meadows). This result suggests

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that cattle are showing adaptations to the major features of these mountainous forests due to

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ferality. Furthermore, these results are the basis for management applications such as predictive

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maps of use by semi-feral livestock.

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KEYWORDS

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Nothofagus forests; beef cattle; semi feral cattle; resource selection probability functions (RSPF);

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GPS tracking; habitat distribution modeling.

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INTRODUCTION

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Cattle (Bos taurus) are an important species for forest dynamics globally, and the originally

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domesticated animals have transgressed to become semi-feral to feral again in many areas

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(Decker et al. 2015, Berteaux and Micol 1992, Hernandez et al. 1999, Baker 1990, Atkinson

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2001, Lazo 1994). Livestock can affect the diversity and structure of plant communities

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(Coughenour 1991, Cingolani et al. 2008) mainly through diet selection (Rook et al. 2004), but

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also due to indirect actions such as changing the distribution of nutrients and compacting the soil

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by trampling (Milchunas et al. 1998, Cingolani et al. 2008). Thus, it is possibly one of the main

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ecosystem engineers for these environments, as has already been demonstrated in grasslands

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(Jones et al. 1997). Although the impacts of cattle are often studied, their habitat use, which

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would allow a comprehensive understanding of the livestock-forest ecological system, has

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generally been overlooked.

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Southern beech forests (i.e., Nothofagus forests) in Patagonia have been used to raise

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cattle since the late 19th century. The traditional management involves extensive cattle ranching

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in the forest. But occasionally and for different reasons, some animals escape the herd,

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establishing feral populations (Atkinson 2001, Veblen et al. 1996). Although feral cattle can be

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found in many areas of the Nothofagus forests in Patagonia (Novillo and Ojeda 2008) and New

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Zealand (Howard 1966, Veblen and Stewart 1982), no studies have been performed on their

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behavior or habitat selection in southern forests. Additionally, many levels of ferality already

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exist depending on contact with human settlements, hunting pressure, isolation time, etc.

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The proportion of time an animal spends in a location to meet their biological demands
defines its habitat use (Beyer et al. 2010). As resources are often heterogeneously distributed,

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individuals need to explore multiple environments to satisfy their needs (Law and Dickman

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1998). Therefore, the selection of habitats is a key feature of behavior and population dynamics

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(Morris 1987). This is especially relevant for large herbivores, because their distribution on the

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landscape can decisively influence the productivity and biodiversity of fields and pastures

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(Bailey and Provenza 2008).

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There are multiple relationships between livestock and their environment but one of the
major ones is the change in the heterogeneity of vegetation produced by foraging (Adler et al.

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2001). In the case of forests, despite having high heterogeneity and frequent cattle presence,

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there is no systematic review of the resulting ecological processes due to the introduction of this

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herbivore worldwide. Furthermore, it is common to have little or no information about the

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activity patterns of semi-wild cattle in general, which makes their management even more

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difficult. Some studies that have been conducted in different forests account for diet selection

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along seasons (Marquardt et al. 2010) or in relation to anthropogenic changes, usually due to

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forest management practices (Roath and Krueger 1982, Kaufmann et al. 2013). In these studies,

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vegetation type and topography were the most common variables to explain habitat selection by

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cattle in forests.

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A variety of technological and statistical tools can be applied to study habitat use. Among

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the first, radiotelemetry and GPS devices allow for accurate and frequent data collection on

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animal location, which facilitates the development of mechanistic models of habitat use and

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movement (Beyer et al. 2010). Therefore, their use is widespread (Johnson et al. 2004, Ungar et

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al. 2005, Cagnacci et al. 2010, Peinetti et al. 2011). In addition to these data collection methods,

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habitat use can be modeled by resource selection functions (RSF) and resource selection

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probability functions (RSPF) that estimate the probability that a given environment has been

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used by an animal (Boyce and McDonald 1999, Beyer et al. 2010). Thus, the RSFs provide a

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framework for developing an ecological theory of habitat use and allow linking landscape

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ecology and population biology (Boyce and McDonald 1999, Boyce 2006). One way to build a

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RSPF with GPS data is to model the intensity of use in sampling units, relating the relative

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frequency of records per unit to the environmental features of such sampling units (Nielson and

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Sawyer 2013).

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The objective of this study was to evaluate the habitat use of semi-feral cattle in southern

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beech forests as no current data or a common methodology exists for this purpose. Furthermore,

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the use of vegetation variables is usual in studies on habitat use by cattle, but the role of

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topography could be of importance especially in mountainous areas, yet is often overlooked.

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Specifically, this study address the following questions: (i) What is the habitat use of semi-feral

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cattle in Nothofagus forests?; and (ii) Are the vegetation variables of the landscape more

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important than topography for habitat use by cattle?. These research questions were addressed by

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using a utilization distribution approach and the construction of a resource selection probability

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function (RSPF) which included topographic and vegetation variables.

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METHODS

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- Study Area

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The study was carried out in the Llodconto Valley, Northwest Patagonia (Rio Negro, Argentina).

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The valley is within the Nahuel Huap National Park, between 4121' and 4127' south latitude,

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and 7131' and 7141' west longitude, at an elevation ranging from 920 to 2000 m.a.s.l.. It is a

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mountainous area with cold-temperate to moist-cold climate with prevailing westerly winds. The

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summers are dry and winters rainy, with a mean monthly temperature ranging from 2.4C in July

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to 12.9C in January. The area is part of the Andean-Patagonian forest and includes a wide

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variety of habitats such as forests, shrublands, meadows, burned forest, riparian and high

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mountain areas. The dominant species is Nothofagus antarctica which is one of the main

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components of both forests and shrublands. Other species such as the Nothofagus pumilio tree,

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the Schinus patagonicus shrub, and the Chusquea coleou bamboo are also abundant. An

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uncertain number of free-ranging cattle inhabit the valley and are traditionally and minimally

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managed by local people only during the summer months.

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- Data collection and explanatory variables

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In order to capture the animals, an enclosure located at the center of the valley was used to herd

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the animals into. Four female adults caught in 2012 and one more caught in 2013 were fitted

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with custom-made GPS collars (GPS Module ZX4120 and Amicus GPS Shiled Antenna,

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Crownhill Associates Ltd.). These GPS collars store data on board and recapture was necessary

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to recover the data. Each collar was set to record location every ten minutes and these records

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were later screened by hour in order to standardize the records and eliminate the unacquired data.

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All capture and handling methods used in this study met the standards of animal welfare

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practices recommended for scientific research (Rollin and Kessel 1998, Gannon and Sikes 2007).

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All research was approved by the proper authorities of the protected area.

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Topographic mapping and spatial statistical analysis were conducted using Quantum

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GIS software (QGIS) version 1.8.0 using a digital elevation model (ASTER GDEM) as the

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data source. Animal locations were overlayed on the digital elevation model and a minimum

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convex polygon (MCP) with a 200 meter buffer was used around the complete set of GPS

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positions to define our study area. One thousand sampling units of 100 meters radii each (i.e.,

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sampling unit area= 0.03km2) were allocated randomly inside this area.

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Habitat types were assigned for each sampling unit by means of a visual classification on

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Google Earth ( Google Inc.), identifying the following vegetation types: forests, shrubland,

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burned areas, meadows, open areas (vegetation cover <25%) and high Andean vegetation. All

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continuous variables were standardized in order to allow comparison between them.

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Corresponding values for elevation, slope, aspect, hill-shade and ruggedness were extracted from

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GIS for each sampling unit. All these were used as predictor variables in the habitat use model.

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An additional variable called "NWness" was created to take into account hillside exposure. The

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rationale for this is based on strong west winds and large thermal amplitudes due to sunlight

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exposure in north facing sites that shape the rigorous weather in these zones. NWness was

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calculated as the cosine of the hillside aspect minus 310 degrees. Thus, this variable accounts for

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the exposure of the slope with a value ranging from 1 (maximum exposure) to -1 (minimum

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exposure).

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Highly correlated variables (r > 0.6) were not used together in the same model. Squared

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elevation and slope were included in the models for the purpose of evaluating preferred ranges of

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use by animals. In order to account for a strong regional climatic pattern, the interaction between

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slope and NWness was also considered since steep and northwest exposed slopes are generally

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arid, with short vegetation and low coverage.

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- Data analysis

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There are several ways to model habitat use. The most widely used include the GLMs and related

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models (Guisan and Zimmermann 2000), but multivariate approaches (Clark et al. 1993, Burke

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et al. 2013), GIS-based habitat suitability models (Osborne et al. 2001, Store and Jokimki

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2003), individual based models (Railsback and Harvey 2002), artificial neural networks

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approaches (zesmi and zesmi 1999), and maximum entropy modeling (Baldwin 2009) also

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exist. Here I applied a utilization distribution approach described by Nielson and Sawyer (2013)

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to develop the RSPF because it accounts for intensity of use among habitats and is unbiased in

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the face of temporally correlated animal location data.

The utilization distribution approach allows the modeling of habitat use by using the

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relative frequency of positions within each sampling unit as an empirical estimator of the use

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distribution (Millspaugh et al. 2006). This relative frequency of positions was used as a response

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variable in a generalized linear model (GLM) with negative binomial distribution that accounts

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for over-dispersed count data. In order to relate the observed counts in a sampling unit with the

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total number of positions recorded for each animal, an offset term was included, thus allowing for

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estimating the probability of use by individual ([i]/total, references below).

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Thus, the RSPF was set as follows:

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yi ~ NegativeBinomial i ,

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ln i = ln total + 0 + 1.elevation i 2 .slopei 3 .ruggedness i 4 .NWness i 5 . forest i ...

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where:

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yi = number of observations at the i-th sampling unit; = relative frequency in each sampling

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unit by animal; = overdispersion parameter; total = number of recorded positions in the whole

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study area by animal; i = sampling unit with associated habitat covariates.

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Several models exploring different combinations of predictor variables were fitted for

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each animal and compared using the Akaike information criterion (AIC). The overdispersion

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parameter of the negative binomial distribution was estimated simultaneously with the models

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using the glm.nb function in R (Venables and Ripley 2002, R Core Team 2013 version 3.0.2). A

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set of ecologically plausible models was chosen to work with all animals based on a stepwise

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regression. The package glmulti (Calcagno and Mazancourt 2010) was used as a tool for quickly

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identifying the best models for each animal. The deviance (Guisan et al. 2000) was used as a

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goodness of fit indicator.

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A population-level model was then developed considering each animal as an

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experimental unit, which reflects the individual nature of resource selection (Marzluff et al.

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2004, Millspaugh et al. 2006). A single set of covariates was selected considering the comparison

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among models and the biological relevance of the variables. The estimated coefficients, variance

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and confidence intervals were calculated for each parameter of this general proposed model.

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The coefficients of the population model were calculated according to:

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1
k = kj
n

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where kj is the estimated coefficient k for each individual j (j=1,2,3,4) and n is the total number

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of animals (n=4).

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The variance of the estimated coefficients for the population model was computed as:

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Var ( k )=

1
2
( ij ij )

n 1 j= 1

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RESULTS

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A total of 2575 position records (an 87% fix rate for the GPS collars) was obtained from 4 of the

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animals. One of the collared animals could not be relocated during the study and therefore was

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not recaptured.

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Based on the fitted models for each animal, the general pattern observed that topographic

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variables were always more important than vegetation. Given the full additive model (not shown)

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as an example, the only meaningful variables for all animals were elevation, aspect and the

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interaction between slope and NWness. In all cases, topographic variables had negative

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coefficients because the animals preferred elevations below the available mean in the landscape

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and avoided hillsides with northwestern exposure and steep slopes. The vegetation variables

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showed different tendencies for each individual and in most cases were not significant (p > 0.05).

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The burned areas and shrubs had a tendency of being avoided by cows 1 and 2 but were

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preferred by cow 3, which also preferred wet meadows. Forest habitat had low coefficients for all

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the animals with values near zero, which showed no tendency to be selected, either in favor or

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against.

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By means of a step-by-step selection procedure over all the models, five plausible

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ecological models were selected for each animal in order to use them as a RSPF for semi-wild

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cattle (Table 1). These models were among the best ten for each animal. It can be seen that all

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models have as variables elevation, some aspect indicator, its interaction with slope, and in some

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cases the vegetation types with greater selection: shrubs and meadows. The coefficients of these

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five models for all the animals are summarized in Appendix 1.

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The most parsimonious models for each animal (Table 2) were significantly different

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from the null model (p<0.01) and explained 46% of the total deviance for cow 1, 37% for cow 2,

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53% for cow 3 and 49% for cow 4. All the animals selected lower elevations than available in

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the landscape. Cow 1 also avoided steep slopes and hillsides with northwest exposure. Cow 2

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also avoided slopes with northwest exposure, and the square of the slope shows that it had a

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range of preferred slopes below the available mean at the site. In addition, it had a tendency to

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avoid shrubs and wet meadows. Cow 3, on the contrary, selected wet meadows, shrub sites and

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steep slopes, avoiding northwest hillsides with steep slopes. Cow 4 used slopes with northwest

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exposure. The confidence intervals of the mentioned variables for each individual do not include

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zero, suggesting good accuracy and inference level (Table 2).

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Models with the lowest AIC were different for each individual (Table 1). However, model

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"D" (i.e. including elevation, NWness:slope, shrubland and meadow) is relatively well-ranked

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for all animals and includes a set of variables that summarize the habitat use patterns observed.

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Therefore, model "D" was chosen to build the final predictive RSPF with all the individuals. This

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general proposed model (i.e. model D, Table 3) shows a common selection for an elevation

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below the average available (Figure 1) and against sites with northwest exposure. These were the

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only variables that allow a strong inference, since their confidence intervals excludes zero.

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Nevertheless, some interesting trends of this population model are noteworthy. Shrubland, for

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example, reduces the probability of use for a given site and the opposite occurs with meadows

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(Table 3). A high variability in resource selection can be observed among individuals. Some

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variables (as the interaction slope:NWness) were selected with the same tendency while others

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such as slope, show high variation between animals (Figure 2).

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DISCUSSION

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Using GPS logs and an approach that takes into account the intensity of use, a RSPF that

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accounts for topography and vegetation was built for semi-wild cattle in the Andean-Patagonian

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forest. This study shows that habitat use by livestock in these forests has a large inter-individual

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variation but also common features that would allow a description of a pattern of use.

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Topographic variables affected the probability of habitat use by cattle to a greater extent than

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vegetation types. In the general proposed model, for instance, a change in a single standard

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deviation unit of elevation (~ 300 meters) resulted in a threefold increased in the probability of

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use (Table 3) and the squared slope has a similar role in some of the alternative models (Table 1).

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This suggests that terrain features have a leading role in defining the use of space by livestock,

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perhaps as an adaptation due to the long history of use and because topography is important in

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these mountainous forests.

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Some common features such as the preferred elevation range and the tendency to prefer

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meadows suggests that it would be possible to describe a population-level home-range for a more

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comprehensive description of habitat use. In this regard, the squared elevation would play an

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important role in defining a preferred elevation range, although there seems to be a high

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variability among individuals. This variability could be modeled by a RSPF with hierarchical

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formulation. Cattle tend to slightly avoid shrubland (Table 3). This may be due to a lower

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proportion of trails in this type of thick vegetation, in contrast to other more open areas or those

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that offer better protection from snow, such as forest. Moreover, these paths are unusable in the

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winter. In this scenario, the availability and network of trails would play an important role in

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defining the population-level home-range of cattle, and therefore, in determining habitat use. At

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the same time, use by livestock has created the paths over time, so there is a reciprocal process

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that shows a type of population memory process.

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The response variable used in this study proved to be appropriate and useful as it

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describes the habitat use of cattle in probabilistic terms. However, the experimental design has

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limitations that must be made explicit. This includes the low sample size of animals. However,

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the different trajectories and spaces used by the sampled individuals suggest that the study was

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able to capture considerable variability in resource use by cattle (Figures 1 and 2) and even

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studies with one GPS-collared animal have accomplished some general features in the behavior

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of a herd (Moritz et al. 2010). Another limitation is the temporal span of the data collection. As

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the data collection period occurred between the months of April and July, it can be assumed that

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this covers a typically cold season (corresponding to the austral autumn and winter). Therefore,

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all models and conclusions of this work should be most reflective of that season.

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A disadvantage of using logistic regressions such as RSPFs is that the model coefficients

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decrease as the availability of a resource increases, and may even change the sign (Beyer et al.

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2010). This could explain some of the inter-individual variation observed in the case of meadow

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use (Figure 2), with animals 1 and 2 both negatively selecting this vegetation, and moving into

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an area where the abundance of meadows is higher in relation to the areas where individuals 3

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and 4 moved (unreported movement data). A similar case is that of the forest variable, which was

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not significant in any model (Table 1) and yet it is a very abundant vegetation type and widely

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used by cattle. To overcome this limitation it would be necessary to develop models that also

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account for animal movement, as habitat selection and movement are not independent processes.

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Methods for estimating movement patterns and habitat selection simultaneously have recently

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been developed (Moorcroft and Barnett 2008, Smouse et al. 2010).

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Comparing the results of this study with others conducted on cattle in forests we find

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interesting relationships in the applied variables and scales. Studies on seasonal diet selection

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(Marquardt et al. 2010) in Bolivian forests for example, show that different functional groups of

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plants are preferred in different seasons. It would be possible to infer species selected by

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livestock throughout the year in the Andean-patagonian forest if direct observations of consumed

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plants was added to this study. Roath and Krueger (1982) also used topographic and vegetation

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variables to study the behavior of cattle in a forested mountainous environment, finding that the

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type of vegetation and water availability determined the degree of use by livestock. These

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apparently opposite results are consistent if we consider the differences between the study areas.

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While Roath and Krueger (1982) worked in a dry, arid vegetation and elevational range of 300

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meters, Llodconto Valley is an area with high water availability, abundant streams, and altitudes

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ranging from 800 to 1900 masl. Kaufmann et al. (2013) found that in silvopastoral systems cattle

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prefer places with forest cover, avoiding places with high slope and low biomass, particularly

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logged sites. A comparison with this study would make an analogy between logged and burned

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areas, both without forest cover. Nevertheless, with the available data we cannot make inferences

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about the type of selection on burned areas so far (Tables 1- 3). It may be noted, however, that

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other studies in Patagonia (Blackhall et al. 2008, De Paz and Raffaele 2013) found a significant

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influence of livestock on vegetation in burned environments, making it clear that cattle forage in

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these areas. Interestingly, Johnson et al. (2004) conducted a study on resource selection by

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caribou at two spatial scales, finding that topographic variables such as elevation and slope are

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more important at a landscape scale, while vegetation variables were more important at the patch

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scale. Something similar could be occurring in this present study, which is closer to the

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landscape scale.

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As mentioned, cattle can increase landscape heterogeneity, especially when the spatial

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pattern of vegetation is already heterogeneous (Adler et al. 2001) as in the case of the currently

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studied forests. This has implications for forests dynamics because although there are studies on

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cattle inhabiting forest (Bartolom et al. 2011, Kauffmann et al. 2013, etc.), there still remains a

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lack of knowledge regarding how forest dynamics are affected by the distribution of wild cattle

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worldwide. Recently, a study conducted in Patagonia developed a model that postulates distance

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to meadows as an estimator of forest use by livestock (Quinteros et al. 2012). While meadows

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are of crucial importance to the Andean-Patagonian forests and Quinteros et al. (2012) make a

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practical contribution in relation to forest management, their model has a spatial limitation, being

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restricted to meadows surroundings. Thus, it is not possible to make inferences at a landscape

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scale and the model fails to consider other relevant variables such as topography, which proved

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to have greater importance in resource selection by cattle (Tables 2-3, Figure 2). Nonetheless, the

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approaches are consistent, because as the distance to meadows increases, the intensity of use by

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livestock decreases (Quinteros et al. 2012), although altitude is also higher, which is predicted by

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the model in the current study.

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There remains a need for tools that enable animal management in agreement with

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conservation management practices of native forests. In Patagonia, for instance, extensive

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livestock farming has been identified as an activity with potential to be carried out in the forest.

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Yet, there have been only some pioneer experiences of silvopastoral handling (Fertig and Guitart

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2006), and current livestock handling is still scarce (Ormaechea et al. 2009). The main

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limitations for management worldwide are a lack of understanding of animal foraging decisions

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and the spatial scale of diet selection (Rook et al. 2004). Although effective management

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depends heavily upon knowledge about the interaction of the animal with its environment, it is

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common to have poor information on the activity patterns of semi-wild cattle (Moyo et al. 2012).

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This current study provides a direct contribution to this knowledge-gap, and direction for future

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research in these systems. As an example, a practical application of this work is the production of

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maps showing intensity of land use by livestock. Similar maps have been developed for other

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regions using equivalent methodologies (Nielsen et al. 2003, Sawyer et al. 2009), and therefore

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offer a management and research tool within protected areas or productive forests.

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In summary, the present study demonstrated that the habitat use by semi-feral cattle in

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southern beech forests has a large inter-individual variability but also similar characteristics

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which allow the description of a pattern of use. Specifically, the results indicate that vegetation

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variables are no more important than topography features. Conversely, habitat selection by cattle

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is mainly affected by topographic variables such as altitude and the combination of slope and

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aspect. These results suggest an adaptation by free-ranging cattle to the major features of the

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landscape of these mountainous forests maybe due to ferality.

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ACKNOWLEDGEMENTS

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Funding for this research was provided by a PhD grant from the CONICET (Argentina). The

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author thanks to Juan Manuel Morales for helpful comments on the manuscript.

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Table 1. Alternative models for free-range cattle habitat use in the study area; the table shows

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Akaike information criterion value (AIC), number of predictor variables (K ) and ranking of

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models relative to each individual (R).

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Models

Animal 1

Animal 2

Animal 3

Animal 4

K AIC AIC R AIC AIC R AIC AIC R AIC AIC R

Elevation + elevation2 +
A slope + slope2 + NWness 5 187.5 0.0

1 619.9 2.4

3 516.0 5.8 2 1813.5 18.1 4

7 188.9 1.4

2 622.7 5.2

5 510.2 0.0 1 1809.2 13.8 3

4 189.5 2.0

3 617.5 0.0

1 528.9 18.7 5 1795.4 0.0 1

6 192.4 4.9

4 619.8 2.3

2 520.5 10.3 3 1795.5 0.1 2

6 196.3 8.8

5 620.8 3.3

4 521.7 11.5 4 1832.1 36.7 5

Model A + shrubland +
B meadow
Elevation +
C NWness*slope
Model C + shrubland +
D meadow
Elevation + slope +
Nwness + forest +
E shrubland + meadow
543
544
545
546
547

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Table 2. Coefficients and 95% confidence intervals of the most parsimonious RSF models for

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selection of topographic and vegetation resources during autumn by free-range cattle in the study

551

area.

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Variables

Intercept

Animal 1

Animal 2

Animal 3

Animal 4

Coef. 95% CIs

Coef. 95% CIs

Coef. 95% CIs

Coef. 95% CIs

-39.91 -67.66 -12.16 -2.73 -4.05 -1.61 -2.22 -3.44 -1.07 -1.59 -2.38 -0.83

Elevation -56.92 -110.64 -3.20 -1.54 -2.90 -0.46

Elevation
Slope

-1.68 -2.82 -0.73

-2.65 -4.21

-1.11 -1.43 -2.79 -0.33 1.45 0.68 2.38 0.73 0.20 1.30

Slope
NWness

-0.26 -1.20 0.48 1.08 0.47 1.76

-2.64 -5.27

-0.00 0.77 0.12 1.46 1.15 -0.01 2.54 2.01 1.29 2.73

Shrubland
Meadow
NW*Slope
553
554
555
556
557
558
559

-3.54 -4.63 -2.56

-0.58 -1.63 0.53

-1.21 -3.46

1.04

4.01 1.61 7.54

-1.42 -2.90 -0.26 1.99 1.38 2.63

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561

Table 3. Population model of resource selection by cattle, with estimate coefficients

562

(standardized variables), SEs and 90% percentile confidence intervals.

563

90% Confidence intervals

Coefficient

564
565
566
567
568
569
570
571

SE

Lower limit

Upper limit

Intercept

-9.368

1.136

-12.409

-6.327

Elevation

-3.016

0.864

-4.776

-1.255

Slope:Nwness

-4.772

1.319

-8.867

-0.677

Slope

-5.958

1.799

-13.576

1.660

NWness

-8.165

1.598

-14.180

-2.151

Shrubland

-0.060

0.795

-1.432

1.545

Meadow

1.521

1.361

-2.839

5.882

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Figure 1. Use vs. Availability of topographic resources elevation and slope.

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576

Figure 2. General proposed model estimates (black) and individual model estimates (grey) with

577

90 % confidence intervals.

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579
580
581
582
583
584
585
586
587
588
589
590
591
592
593
594

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597

Figure 1

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603
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607
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609
610
611
612
613
614
615
616
617

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Figure 2

621
622
623
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628
629
630
631
632
633
634
635
636
637
638
639
640

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Appendix: The models and their coefficients for all animals.

643

############################################################

644

1) Description of the models

645

2) Coefficients of all tested models and animals

646

############################################################

647

1- Description of the models

648

yi ~ NegativeBinomial i ,

649

ln i = ln total + 0 + j .variablei

650

where:

651

y = number of observations per sampling unit; = relative frequency in each sampling unit by animal; =

652

overdispersion parameter; total = number of recorded positions in whole study area by animal; i = sampling unit

653

with associated habitat covariates; j = number of variables in each model.

654

655
656
657
658
659

Model

Description of variables involved in each model

elevation + sqr.elev + slope + sqr.slope + NWness

elevation + sqr.elev + slope + sqr.slope + NWness + shrubland + meadow

elevation + slope + NWness * slope

elevation + NWness * slope + shrubland + meadow

elevation + slope + NWness + forest + shrubland + meadow

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661
662

2- Coefficients of all tested models and animals*

663

* Coefficient with this symbol are significant at a level of 95%

664

# Animal 1
Model A

Model B

Model C

Model D

Model E

Intercept

-72.577*

-75.991*

-11.633*

-11.422*

-10.516*

Elevation

-130.564*

-137.462*

-5.748*

-5.718*

-5.394*

Elevation2

-62.145*

-65.576*

Slope

-2.343

-2.657

-3.063*

-3.051*

-2.536*

Slope2

-0.101

-0.236

NWness

-0.060

-0.043

-2.852*

-2.621*

-0.010

Shrubland

-0.259

-0.284

-0.252

Meadow

-1.681

-1.127

-1.218

Forest

0.174

NWness*Slope

-2.279*

-2.117*

Theta

0.212

0.238

0.164

0.168

0.160

Theta SE

0.098

0.113

0.067

0.069

0.067

Model A

Model B

Model C

Model D

Model E

Intercept

-3.174*

-2.686*

-3.614*

-3.272*

-3.126*

Elevation

-1.577*

-1.585*

-1.967*

-1.937*

-1.572*

665
666

# Animal 2

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0.127
Elevation2

-0.078

Slope

-1.515*

-1.442*

Slope2

-0.231

-0.252

NWness

0.650*

0.783*

-1.334*

-1.227*

-1.167*

1.039*

1.312*

0.784

Shrubland

-0.627

-0.685

-0.275

Meadow

-0.203

-0.232

0.105

Forest

0.387

NWness*Slope

0.403

0.516

Theta

0.062

0.063

0.063

0.065

0.064

Theta SE

0.011

0.011

0.011

0.012

0.012

Model A

Model B

Model C

Model D

Model E

Intercept

-2.834*

-2.683*

-1.943*

-2.264*

-2.939*

Elevation

-1.512*

-1.636*

-1.713*

-1.698*

-1.711*

Elevation2

-1.288*

-1.597*

Slope

1.652*

1.969*

2.344*

3.010*

2.927*

Slope2

1.196*

1.043*

NWness

0.874*

1.444*

0.233

1.034*

0.994*

667
668

# Animal 3

Shrubland

-0.672

-0.712

-0.202

Meadow

2.929*

3.937*

4.166*

Forest

1.340

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NWness*Slope

-0.161

-0.873*

Theta

0.040

0.044

0.031

0.037

0.036

Theta SE

0.008

0.008

0.006

0.007

0.007

Model A

Model B

Model C

Model D

Model E

Intercept

0.073

-1.003*

-1.594*

-1.817*

0.230

Elevation

-3.173*

-3.131*

-3.545*

-3.420*

-2.639*

Elevation2

-1.273*

-0.782*

Slope

1.471*

1.591*

0.739*

0.783*

1.364*

Slope2

0.714*

0.830*

NWness

0.214

0.132

2.012*

1.740*

-0.177

669
670

# Animal 4

Shrubland

1.120*

0.634

-0.053

Meadow

0.212

0.794

0.764

Forest

-0.647

NWness*Slope

671
672
673

1.996*

1.872*

Theta

0.072

0.075

0.078

0.079

0.068

Theta SE

0.007

0.007

0.007

0.008

0.006