**2. Basic concepts**

282 Modern Telemetry

to remotely locate wild species and obtain ecological, behavioural and physiological data. It was in the 1960s, when telemetry, specifically radio-telemetry, was first used to study terrestrial wildlife (Craighhead, 1982; Hebblewhite & Haydon, 2010). Since then, wildlife telemetry has contributed significantly to our understanding of fundamental ecological and behavioural processes of many animal species (e.g., Johnson et al., 2006). Advances in wildlife telemetry have made it possible to acquire detailed data on animal' space use, including habitat selection, home-range size, movement metrics as well as migration timing and routes. Since many wildlife species are secretive and difficult to observe, telemetry has

Human attitudes vary worldwide towards mammalian carnivore species. The overlap in space-use with human populations results in competition for habitats and resources which is at the heart of most of the conflicts between mammalian carnivore species, especially canids, and humans (Sillero-Zubiri & Switzer, 2004). For instance, canids tend to prey upon a range of livestock, game stock and threatened wildlife, and some of the large-bodied size species may also attack, and on rare occasions, deadly harm humans. Human–carnivore conflicts are among the major causes of population decline in many species (Treves & Karanth, 2003) and can be particularly controversial when the resources concerned have economic value (e.g., livestock) and the carnivore species involved have a high conservation profile (Graham et al., 2005). This is usually the case of large-bodied size carnivore species that usually have extensive space requirements, low reproductive rates and are persecuted by humans (Matthiae & Stearns, 1981). But, while these carnivore species are declining globally, others have not only managed to survive, but to become abundant. Many mediumbodied size generalist carnivores have been able to expand their geographic ranges because they are capable of using human-use areas and anthropogenic resources (Harris & Smith, 1987; Prange et al., 2004). For instance, red foxes (*Vulpes vulpes*) and coyotes *(Canis latrans)*  may occur not only in rural areas, but also in suburban and occasionally more densely populated urban areas (Atkinson & Shackleton, 1991; Gibeau, 1998; Grinder & Krausman,

In this study, we will consider the use of telemetry data to investigate habitat selection and home-range patterns of two mammalian carnivore species. Although both species occur in human-dominated landscapes, their interactions with humans are very distinct, resulting in very different abundance levels and conservation status. However, in both cases, investigations of habitat selection and home-range patterns are fundamental initial steps in the management of their populations. In the following Section 2, we will briefly define key concepts that will be used in this chapter. In Case Study 1 (Section 3), we will use telemetry data to investigate habitat selection and home range of the red fox in Prince Edward Island, Canada. In this case, the species is benefiting from its interaction with human populations. Our objective for Case Study 1 is to show how telemetry data can be used to elucidate the effects of fox-feeding (anthropogenic food resources provided to foxes by humans) on habitat selection and home-range patterns. In Case Study 2 (Section 4), telemetry data will be used to investigate habitat selection and home range patterns of the African wild dog (*Lycaon pictus*) in Mkhuze Game Reserve, South Africa. Contrary to the red fox, the interaction with humans has had detrimental effects on African wild-dog populations in South Africa and other parts of Africa. Our objective for Case Study 2 is to illustrate the use of telemetry data to assess the success of the establishment part of a reintroduction program of a endangered or threatened carnivore species. For purposes of comparison, we have tried to use the same approaches, methods and data-analysis procedures for both case studies

become a valuable tool to learn more about their respective life-histories.

2001; Lewis et al., 1999).

*Habitat* refers to a distinctive set of physical environmental factors that a species uses for its survival and reproduction (Block & Brennan, 1993). The semantic and empirical distinctions between the terms habitat *use* and habitat *selection* are often unclear (Hall et al., 1997). Habitat *selection* carries a connotation of understanding complex behavioral and environmental processes that habitat *use* does not; habitat-use patterns are the end result of habitat-selection processes. *Use* of habitat is the way in which an individual or species exploits habitats to meet its life history needs (Block & Brennan, 1993). The study of habitatuse patterns describes the actual distribution of individuals across habitat types (Hutto, 1985). *Selection* of habitat is the process by which an animal actually chooses habitat (Johnston, 1980). In other words, habitat *selection* refers to a hierarchical process of behavioral responses that may result in the disproportionate *use* of habitats to influence survival and fitness of individuals (Block & Brennan, 1993; Hutto, 1985). *Use* is considered selective if habitat is used disproportionately compared with its *availability*, the latter being the amount of that habitat *accessible* to the animal. In field studies, however, where the *availability* of habitat is variable, habitat selection is the use of habitat relative to its availability in the environment and is conditional on the availability of all habitats to the animal. It is important to distinguish between the terms *accessible* and *available* because it may be possible that certain habitats, within a given landscape, are available to an animal (or species), but they may not be accessible. Use-availability studies inherently assume that study animals have free and equal access to all habitats considered to be available, implying that at any given moment each studied animal should be able to use any available habitat (Garshelis, 2000). This assumption may stand if use and availability are measured for each animal individually, but it may be violated when data from different animals are pooled together and the available habitat is considered to be same for all when it is not the case. For instance, use and availability may be considered to be the same when all animals move as a pack or in cases when there is significant overlap in their home ranges. However, differences between use and availability may occur when not all animals have the same habitat types within their home ranges (Garshelis, 2000). Differences between use and availability may also occur when not all animals have free or equal access to all areas within their home ranges.

Habitat-selection scales are often assumed to be a function of home-range sizes (e.g., Chamberlain et al., 2003; McLoughlin et al., 2002, 2004; Rettie & Messier, 2000). Thus, an important concept associated with habitat selection is home range. Burt (1943) first defined home range as the area traversed by an individual when performing normal activities such as foraging, mating and caring for young. However, this definition has been challenged because the word "normal" is difficult to interpret and lacks a temporal component (Cavallini, 1996; White & Garrott, 1990). A less ambiguous, and more popular, definition of the home range of an animal is the limited area within which it can be found during a specified time period (Harris et al., 1990; Kernohan et al., 2001). According to this definition,

Use of Telemetry Data to Investigate

by vehicles.

**3.1 Study site** 

population dynamics of red foxes in Prince Edward Island.

Home Range and Habitat Selection in Mammalian Carnivores 285

characterizing human-altered landscapes (Ables, 2009; Catling & Burt, 1995; Lloyd, 1975). In Prince Edward Island (Canada), the red fox was the largest mammalian carnivore species until the arrival of coyotes during the early 80s. Although no studies have assessed the abundance of red foxes on Prince Edward Island, it is commonly known that they occur throughout the whole province, including urban areas. Regardless of its widespread distribution, little is known about the home range, habitat selection, behaviour and

The red fox is an opportunistic species with a diverse diet that has allowed it to survive in natural and human-altered landscapes (Dell'Arte et al., 2007). Eating habits of red foxes vary, but normally they include entire mice, voles, birds and rabbits. However, the importance of each prey varies depending on habitat type, regional prey availability and anti-predator behavior (Dell'Arte et al., 2007; MacDonald, 1977). Additionally, studies have found that anthropogenic food items can sometimes play an important role in the diet of red foxes (e.g., Contesse et al., 2004; Newsome et al,. 2010). Another relevant characteristic of red foxes is that their interactions with humans can take different forms. In areas where they are potential carriers of the rabies virus, red foxes are considered nuisance animals and are usually subject to population control even though these operations are generally unsuccessful (Smith & Harris, 1991). In Prince Edward Island, especially within and near Prince Edward Island National Park, red foxes are considered charismatic animals and many residents and tourists feed them throughout the year. Red foxes are fed by humans inhabiting houses or cottages located near the park, as well as by tourists on roadsides. Little is known about the consequences of this activity on red fox population occurring within Prince Edward Island National Park. However, it has been postulated that because humans feed these animals in the park, red foxes tend to select road and human-use habitats, rather than natural habitats for foraging, thus increasing the probability of being accidentally killed

In this study, we determined habitat selection and home range of red foxes occurring in Prince Edward Island National Park using radio-telemetry data. Specifically, we examined the use of radio-telemetry data to elucidate the importance of human-use areas and fox-

Prince Edward Island is situated in the Gulf of the St. Lawrence and encompasses an area of about 5,660 km2 (Weighs, 1995). Prior to the 17th century, Prince Edward Island was covered by tree species characteristic of the Acadian Forest region, such as sugar maple (*Acer saccharum*), yellow birch (*Betula alleghaniensis*) and beech (*Fagus grandifolia*) (Round Table on Resource Land Use and Stewardship, 1997). Since the arrival of European colonizers about three centuries ago, anthropogenic activities such as urbanization, forestry, and agriculture have altered the natural habitats of the island. Although the peak of deforestation occurred in the early 1900s, it is the intensive exploitation for agriculture that have occurred during the last century that have resulted in most of the major transformation of the natural habitats of Prince Edward Island (Johnston, 2000). Currently, the forests of Prince Edward Island are composed of species such as white spruce (*Picea glauca*), balsam fir (*Abies balsamea*), and trembling aspen (*Populus tremuloides*) (Round Table on Resource Land Use and Stewardship, 1997). It is possible to assume that these changes in the structure and composition of vegetation have been followed by significant alterations in the distribution and abundance of natural resources for

feeding in habitat selection and home range patterns of red foxes.

mammalian species occurring in Prince Edward Island.

a home range can be flexible, varying with season and overlapping with conspecifics (Harris et al., 1990), making the concept of home range particular useful for habitat selection studies. In contrast, a territory, a term commonly used interchangeably with home range, is defined as an area that is occupied by an individual or group to the exclusion of other animals of the same species (Börger et al., 2008; Burt, 1943; Mech, 1970). Animals may or may not be territorial, but will still have a home range.

Quantifying an animal's home-range size and shape allows researchers to gain information on foraging behaviour and inter- and intraspecific interactions (Harris et al., 1990). It is also useful for investigating animal-habitat relationships such as habitat selection (e.g., Johnson, 1980). For instance, Johnson (1980) proposed a habitat selection classification system that involves the notion of home range. Johnson's (1980) classification system is based on a hierarchical order of selection: 1st-order selection of a species' geographic range from the global pool, 2nd-order selection of home ranges from the geographic range, 3rd-order selection of habitat within home ranges (e.g., *core areas*), and 4th-order selection of structures, variables or conditions within habitats. The term *core area* refers to those areas within the home range where individuals are found with greater probability (Börger et al., 2008; Kaufmann, 1962; White & Garrott, 1990). Thus, core areas are locations of concentrated use within home ranges (Kaufmann, 1962) that contain important resources such as den sites and quality foraging areas (Ewer, 1968). Although core areas may contain similar landscape elements as the whole home-range area, the importance of a given habitat type can vary between "core areas" and "home-range size". While home ranges of different animals have been found to overlap (e.g., Kolb, 1986; Lovari et al., 1994), overlap of the core areas does not commonly occur (Samuel et al., 1985). Thus, the identification of core areas is important when studying intraspecific interactions or when investigating animal-habitat relationships (Samuel et al., 1985).

Another important factor that must be taken into consideration when investigating habitat selection patterns is that many resources used by wild species occur heterogeneously across the landscape, linking the concept of habitat selection with the ideas of space-use and scale. For instance, the process of habitat selection in mammalian carnivores has increasingly been studied as a hierarchical, multi-scale process, in which selection of habitat features is accomplished at progressively smaller scales (Orians & Wittenberger, 1991; Rettie & Messier, 2000; Schaefer & Messier, 1995). An organism selects a home range in which to live, and then it makes decisions about the use of different habitats within this home range in which fundamental activities such as foraging will be performed (Johnson, 1980). Rettie & Messier (2000) proposed that animal selection patterns that are governed by an avoidance of factors that tend to limit individual fitness dominate at the larger spatial scales, while less important limiting factors influence habitat selection patterns at smaller spatial scales. If Rettie & Messier's (2000) hypothesis holds true, then processes occurring at the larger spatial scales exert the most influence on species habitat selection. One may then predict that habitat quantified at the largest spatial scales would explain or describe the most variability in species habitat selection.

#### **3. Case study 1: The red fox**

The red fox is a generalist species that can occur in a variety of habitats, including forest, tundra, agricultural land, desert and urban areas. Several studies, however, have shown that this species tends to be more abundant in mixed mosaic habitats such as those characterizing human-altered landscapes (Ables, 2009; Catling & Burt, 1995; Lloyd, 1975). In Prince Edward Island (Canada), the red fox was the largest mammalian carnivore species until the arrival of coyotes during the early 80s. Although no studies have assessed the abundance of red foxes on Prince Edward Island, it is commonly known that they occur throughout the whole province, including urban areas. Regardless of its widespread distribution, little is known about the home range, habitat selection, behaviour and population dynamics of red foxes in Prince Edward Island.

The red fox is an opportunistic species with a diverse diet that has allowed it to survive in natural and human-altered landscapes (Dell'Arte et al., 2007). Eating habits of red foxes vary, but normally they include entire mice, voles, birds and rabbits. However, the importance of each prey varies depending on habitat type, regional prey availability and anti-predator behavior (Dell'Arte et al., 2007; MacDonald, 1977). Additionally, studies have found that anthropogenic food items can sometimes play an important role in the diet of red foxes (e.g., Contesse et al., 2004; Newsome et al,. 2010). Another relevant characteristic of red foxes is that their interactions with humans can take different forms. In areas where they are potential carriers of the rabies virus, red foxes are considered nuisance animals and are usually subject to population control even though these operations are generally unsuccessful (Smith & Harris, 1991). In Prince Edward Island, especially within and near Prince Edward Island National Park, red foxes are considered charismatic animals and many residents and tourists feed them throughout the year. Red foxes are fed by humans inhabiting houses or cottages located near the park, as well as by tourists on roadsides. Little is known about the consequences of this activity on red fox population occurring within Prince Edward Island National Park. However, it has been postulated that because humans feed these animals in the park, red foxes tend to select road and human-use habitats, rather than natural habitats for foraging, thus increasing the probability of being accidentally killed by vehicles.

In this study, we determined habitat selection and home range of red foxes occurring in Prince Edward Island National Park using radio-telemetry data. Specifically, we examined the use of radio-telemetry data to elucidate the importance of human-use areas and foxfeeding in habitat selection and home range patterns of red foxes.

#### **3.1 Study site**

284 Modern Telemetry

a home range can be flexible, varying with season and overlapping with conspecifics (Harris et al., 1990), making the concept of home range particular useful for habitat selection studies. In contrast, a territory, a term commonly used interchangeably with home range, is defined as an area that is occupied by an individual or group to the exclusion of other animals of the same species (Börger et al., 2008; Burt, 1943; Mech, 1970). Animals may or may not be

Quantifying an animal's home-range size and shape allows researchers to gain information on foraging behaviour and inter- and intraspecific interactions (Harris et al., 1990). It is also useful for investigating animal-habitat relationships such as habitat selection (e.g., Johnson, 1980). For instance, Johnson (1980) proposed a habitat selection classification system that involves the notion of home range. Johnson's (1980) classification system is based on a hierarchical order of selection: 1st-order selection of a species' geographic range from the global pool, 2nd-order selection of home ranges from the geographic range, 3rd-order selection of habitat within home ranges (e.g., *core areas*), and 4th-order selection of structures, variables or conditions within habitats. The term *core area* refers to those areas within the home range where individuals are found with greater probability (Börger et al., 2008; Kaufmann, 1962; White & Garrott, 1990). Thus, core areas are locations of concentrated use within home ranges (Kaufmann, 1962) that contain important resources such as den sites and quality foraging areas (Ewer, 1968). Although core areas may contain similar landscape elements as the whole home-range area, the importance of a given habitat type can vary between "core areas" and "home-range size". While home ranges of different animals have been found to overlap (e.g., Kolb, 1986; Lovari et al., 1994), overlap of the core areas does not commonly occur (Samuel et al., 1985). Thus, the identification of core areas is important when studying intraspecific interactions or when investigating animal-habitat relationships

Another important factor that must be taken into consideration when investigating habitat selection patterns is that many resources used by wild species occur heterogeneously across the landscape, linking the concept of habitat selection with the ideas of space-use and scale. For instance, the process of habitat selection in mammalian carnivores has increasingly been studied as a hierarchical, multi-scale process, in which selection of habitat features is accomplished at progressively smaller scales (Orians & Wittenberger, 1991; Rettie & Messier, 2000; Schaefer & Messier, 1995). An organism selects a home range in which to live, and then it makes decisions about the use of different habitats within this home range in which fundamental activities such as foraging will be performed (Johnson, 1980). Rettie & Messier (2000) proposed that animal selection patterns that are governed by an avoidance of factors that tend to limit individual fitness dominate at the larger spatial scales, while less important limiting factors influence habitat selection patterns at smaller spatial scales. If Rettie & Messier's (2000) hypothesis holds true, then processes occurring at the larger spatial scales exert the most influence on species habitat selection. One may then predict that habitat quantified at the largest spatial scales would explain or describe the most variability

The red fox is a generalist species that can occur in a variety of habitats, including forest, tundra, agricultural land, desert and urban areas. Several studies, however, have shown that this species tends to be more abundant in mixed mosaic habitats such as those

territorial, but will still have a home range.

(Samuel et al., 1985).

in species habitat selection.

**3. Case study 1: The red fox** 

Prince Edward Island is situated in the Gulf of the St. Lawrence and encompasses an area of about 5,660 km2 (Weighs, 1995). Prior to the 17th century, Prince Edward Island was covered by tree species characteristic of the Acadian Forest region, such as sugar maple (*Acer saccharum*), yellow birch (*Betula alleghaniensis*) and beech (*Fagus grandifolia*) (Round Table on Resource Land Use and Stewardship, 1997). Since the arrival of European colonizers about three centuries ago, anthropogenic activities such as urbanization, forestry, and agriculture have altered the natural habitats of the island. Although the peak of deforestation occurred in the early 1900s, it is the intensive exploitation for agriculture that have occurred during the last century that have resulted in most of the major transformation of the natural habitats of Prince Edward Island (Johnston, 2000). Currently, the forests of Prince Edward Island are composed of species such as white spruce (*Picea glauca*), balsam fir (*Abies balsamea*), and trembling aspen (*Populus tremuloides*) (Round Table on Resource Land Use and Stewardship, 1997). It is possible to assume that these changes in the structure and composition of vegetation have been followed by significant alterations in the distribution and abundance of natural resources for mammalian species occurring in Prince Edward Island.

Use of Telemetry Data to Investigate

this study.

fields, marshes, and shrubs.

Home Range and Habitat Selection in Mammalian Carnivores 287

recorded 2-3 azimuths within 15-min intervals over a period of 1 hr. During the active periods (late afternoon and early evening), simultaneous triangulation was performed by 2-3 observers within 15-min intervals over a period of 1 hr. In addition, we also conducted 8-hr intensive (1 location every 15 min) telemetry sessions (sequential locations) twice per month (one in the afternoon: 13:00 to 21:00 and one during the night: 21:00 p.m. to 05:00 a.m.). These telemetry sessions were subjected to weather conditions, thus sometimes they were shorter than 8 hr. Tracking was done on foot using hand-held Yagi antennas and portable receivers (R-1000; Communications Specialists, Inc.). Triangulation angles were maintained between 30° and 150° (Gese, 2001). About 50% of radio-tracking locations were taken with the animal in view of the observers. Bearings were plotted immediately using LOAS 2.1 (Ecological Software Solutions 2003) to determine the accuracy of the locations. For the purposes of this study, we have not looked for statistical independence of locations (e.g., Swihart & Slade, 1985), but rather for their biological independence, using a minimum time interval between successive locations long enough to allow any radio-collared animal to cross entirely its home range. Repeated observations on the same individuals, as is the case with radio-telemetry locations, are often assumed to give rise to constant within-group correlation structures (i.e., lack of independence in the data). Although some ecologists and wildlife managers think that correlation structures represent a problem for the analysis and interpretation of wildlife telemetry data (Hansteen et al., 1997), others suggest that much can be learned by studying the causes and consequences of correlation structures in telemetry data. If a large time interval between successive locations is possible and/or the calculation of home-range size is the main goal of the study, strict adherence to the collection of non-autocorrelated data may be necessary. However, it may be difficult to translate autocorrelated data into an independent form and still retain a sample size that is adequate for the home-range size to reach an asymptote. Thus, the goals of most home-range or habitat selection studies require the collection of data which are dependent to some degree. This was the case in

Habitat variables were determined using 2000 Prince Edward Island aerial photographs (1: 17500; Prince Edward Island Department of Agriculture and Forestry) of the study site, and complemented with field observations to update any land-cover changes. ArcView GIS (version 3.3; Environmental Systems Research Institute, Inc., ESRI) was used to map animal locations obtained from radio-telemetry monitoring sessions and to assign to each location a habitat cover-type. Habitat was classified into 1 of 10 cover-types: agriculture (hay fields and pasture), forest (mature white spruce and hardwood forest), water or aquatic systems (frozen during winter; including ponds and saltwater bodies), dunes, beaches, roads (mostly paved), human-use areas (residential areas, recreational areas, parking lots), abandoned

Habitat selection was examined by comparing use and availability of habitat types within the study area using the Neu Method (Neu et al., 1974). The Neu method is a straightforward application of the χ2 goodness-of-fit test, and is usually used to compare observed counts of animals in each habitat with the counts expected if habitats were used in proportion to their availability. The method involves the calculation of confidence intervals (Bonferroni Z-statistic) around the expected proportions to determine whether the observed proportion of usage in each habitat is significantly different from expected. The usage of a particular habitat type was defined as the ratio between animal locations in each habitat type and the total number of locations recorded in the study area. Expected

The study site selected for this study was located in Stanhope (543 ha; 46°25'E; 63°06'N), Prince Edward Island National Park (Fig. 1). Although situated within the park, Stanhope is surrounded by private houses and cottages, many of them inhabited throughout the whole year. It encompasses forest patches, marshes, shrubs, ponds, dunes, beaches, roads, agricultural fields, and human-use areas (about 0.30 human dwellings per ha). Although we do not possess quantitative data regarding fox-feeding intensity in Stanhope, anecdotal information and personal observations clearly indicate that fox-feeding is a common activity in Stanhope, with residents setting out food for foxes throughout the whole year and many tourists feeding these animals during the touristic season (i.e., summer).

Fig. 1. Map of Prince Edward Island (Canada) in relation to Canada indicating the location of the study site (Stanhope) with a star. The scale-bar refers to Prince Edward Island only.
