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Draft

Space use of a reintroduced population of Capra pyrenaica

in a protected natural area

Journal: Canadian Journal of Zoology

Manuscript ID cjz-2015-0166.R2

Manuscript Type: Article

Date Submitted by the Author: 15-Dec-2015

Complete List of Authors: Refoyo Román, Pablo; Complutense University of Madrid, Zoology and Physical Antropology Olmedo, Cristina; Complutense University of Madrid, Zoology and Physical Antropology Muñoz, Benito; Complutense University of Madrid, Zoology and Physical Antropology

Keyword: Capra pyrenaica, Iberian Ibex, density-dependent, Reintroduction, National Park, Spain

https://mc06.manuscriptcentral.com/cjz-pubs

Canadian Journal of Zoology

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1

Space use of a reintroduced population of Capra 1

pyrenaica in a protected natural area 2

P. Refoyo (Corresponding author); C. Olmedo and B. Muñoz. 3

4

P. Refoyo. Department of Zoology and Physical Anthropology, Complutense University of Madrid. C/ José Antonio 5

Novais, 12, E-28040 Madrid. PHONE: +34 91 394 50 31 FAX: 34 91 394 49 47 [email protected] (Corresponding 6

author) 7

C. Olmedo. Department of Zoology and Physical Anthropology, Complutense University of Madrid. C/ José Antonio 8

Novais, 12, E-28040 Madrid. [email protected] 9

B. Muñoz. Department of Zoology and Physical Anthropology, Complutense University of Madrid. C/ José Antonio 10

Novais, 12, E-28040 Madrid. [email protected] 11

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Space use of a reintroduced population of Capra pyrenaica in a protected natural area 13

P. Refoyo (Corresponding author); C. Olmedo and B. Muñoz. 14

ABSTRACT 15

In Europe, wild ungulates have undergone major expansion and population growth during recent decades. In certain 16

cases, the high density achieved by these populations has led to excessive pressure on the environment, which 17

eventually becomes a limiting factor for the population itself. One of these reintroductions was performed with the 18

Iberian ibex (Capra pyrenaica (Schinz, 1838)) in the National Park of the Sierra de Guadarrama (Spain). This 19

reintroduced population was monitored during six field seasons (2000, 2003, 2005, 2007, 2010, and 2014) by direct 20

observation of the animals along transects using the distance sampling method to determine the degree of expansion 21

over the years and the use of different habitats according to different seasons. The abundances obtained for each field 22

season showed a significant increase from 4.16 ind./km to 8.65 ind./km, showing a linear relationship between the 23

abundance and the extent of the area occupied by the species. We observed that differences between habitat 24

availability and use were significant for all seasons.. Our data can be used as an example of the process of colonisation 25

of a population of wild ungulates and their impact on vegetation in order to better manage future reintroductions. 26

27

Keywords: Capra pyrenaica, Iberian Ibex, density-dependent, Reintroduction, National Park, Spain. 28

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INTRODUCTION 30

The introduction or reintroduction of ungulates is closely linked to human activities and movements (Christie and 31

Andrews 1966), Some of these reintroductions have been successful, as the case of the Oryx (Oryx sp.) In Oman or 32

ibex ((Capra ibex) in the Alps or Capra pyrenaica in Peneda-Gerês National Park in northern Portugal (Moço et al., 33

2006), however others have failed as the attempts to introduce reindeer (Rangifer tarandus) in Matthews Island (Bering 34

Sea) (Soriguer et al., 1998). In all cases, these reintroductions should be done as suggested by international 35

organizations (IUCN 1998) that highlights the importance of theparameter of parameters of the population evolution 36

(Converse et al. 2013) and their relationship with the environment (Chapman 1928; Odum 1986). 37

This study goes into detail about the dispersion and use of space of a wild ungulate’s reintroduced population in a 38

Natural Protected Area. 39

40

In Europe, wild ungulates have undergone major expansion and population growth in recent decades, mainly going from 41

marginal areas to occupying a large part of the territory (Apollonio et al. 2010). There have been numerous 42

reintroductions of ungulates in protected natural areas (Geremia et al. 2011) that have enabled better protection of 43

these populations. 44

45

However, these protected natural spaces become restricted areas from where individuals cannot leave without 46

increasing the risk of their survival. This situation leads to a high concentration of ungulates in very limited areas and 47

causes high pressure on vegetation, which becomes a limiting factor affecting the dynamics of the population (Gaillard 48

et al. 2000; Bonenfant et al. 2009). On the other hand, a high density has significant costs for the population (Côté et al. 49

1995; Loehle 1995; Stillman et al. 1997; Krause et al. 2002; Fortin et al. 2009), and it can have important implications in 50

terms of social behaviour (Bateman et al. 2012), survival, and reproduction (Gaillard et al. 1998, 2000; Eberhardt 2002; 51

Festa-Bianchet et al. 2003). 52

53

In our case, the Iberian ibex (Capra pyrenaica (Schinz, 1838)) was distributed in the major mountain ranges of the 54

middle, eastern and southern of the Peninsula until the late nineteenth century. Excessive pressure exerted by men 55

produced its disappearance in much of the territory (Cabrera 1914; Perez et al. 2002). 56

57

The reintroduction of the Iberian ibex in Sierra de Guadarrama National Park began in 1990 with specimens of Capra 58

pyrenaica victoriae Cabrera, 1911 subspecies from a preserve adjacent to the National Hunting Reserve of Gredos and 59

Las Batuecas National Hunting Reserve. A total of 67 specimens (41 females and 26 males) were reintroduced by the 60

time it was finalized in 1992. The average age of the population was around 5 years, and the sex ratio of the introduced 61

specimens favored females at a proportion close to 1.6:1 (41 females and 26 males). However, among juveniles (under 62

5 years), the sex ratio favored males 1:1.4. On the contrary, there was a predominance of females in the individuals 63

older than 5 years in a ratio close to 2:1 (Refoyo et al., 2014). These ratios were in accordance with those used 63 for 64

other ibex reintroductions (Girard et al. 1998). 65

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Emigration and range expansion have been documented in several large ungulate populations when forage quantity or 67

quality has decreased due to density-dependent resource consumption (Lemke et al. 1998; Aanes et al. 2000; Larter et 68

al. 2000; Ferguson et al. 2001; Amarasekare 2004). There are also many studies of modern animal movement relative 69

to partial migration (Hebblewhite and Merrill 2009; Rivrud et al. 2010); however, most of these studies have focused on 70

natural populations, and few have bothered to analyse the process of land occupation of a reintroduced population, as 71

in our case. 72

73

The relationship between habitat selection and the abundance and distribution of species in a particular area has also 74

been widely discussed by several authors (Johnson 1980; Rosenzweig 1981) and constitutes an integral part of 75

effective management and conservation (Boyce and McDonald 1999). Habitat selection occurs over large areas when 76

we consider higher order scales of selection (Johnson 1980; Kittle et al. 2008; Peters et al. 2013), but also very small 77

spatial scales (e.g. warblers in a tree: MacArthur 1958). Similarly, habitat selection can vary over time (e.g. seasons) 78

(Singer 1979; Jenkins and Wright 1988; Macandza et al. 2012) as the forage base is progressively depleted (Brown and 79

Rosenzweig 1986; Van Beest et al. 2010). Nevertheless, there have not been many studies focusing on the effective 80

impact on the core area considering the seasonal movements of the species. 81

82

The aim of this work was to establish changes of the occupied area of the reintroduced Iberian ibex during settlement 83

and colonisation. Our goals were: (1) to determine if there has been an increase in the area of distribution of the species 84

during the settlement process, (2) to determine whether there are relationships between the area of occupation and the 85

detected population increase during the settlement process, (3) to establish if the occupation of the area by ibex differs 86

seasonally and to determine if the pressure on the different plant communities in the park continues throughout the year, 87

and (4) to confirm if there is habitat selection regardless of availability in each season. 88

89

Our data can be used as an example of the process of colonisation of a population of wild ungulates. 90

91

MATERIALS AND METHODS 92

Study area 93

We monitored Iberian ibex in the National Park of the Sierra de Guadarrama. This National Park (Figure 1) has an area 94

of 33.960 ha and it is located in the center of the Iberian peninsula, in the eastern part of the Central System. The area 95

shows a marked difference in altitude (between 2.200 and 1.100 m), alternating between very steep rocky areas (“Las 96

Pedrizas”) and areas of gentle topography. In general, they are very old rocks (from Paleozoic and Mesozoic) such 97

asgneisses, marbles and schists. Sierra de Guadarrama’s climate is typically continental with significant temperature 98

variations between seasons and very dry summers. The average temperature of the coldest month is -0.7°C and 16.4°C 99

in the warmest month, (January and July respectively). The annual rainfall is 1,325 mm with a high seasonal variation, 100

being July (24.9 mm) and November (176.5 mm) the driest months On average, the days of snowfall are 78 and 142 the 101

frost days a year.. The vegetation growing period is 5 months, although is less in the peaks above 2,000 meters .. Its 102

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103

104

105

106

107

108

109

110

111

112

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vegetation includes shrubs (Cytisus purgans, Juniperus communis nana) and grassland (Festuca indigesta, Nardus

stricta, Festuca rubra) in highland areas, areas of Mediterranean shrubs (Cistus ladanifer, Rosmarinus officinalis,

Thymus vulgaris, Lavandula stoechas) in steeply sloped areas, and forests (Quercus ilex, Q. pyrenaica, Pinus spp.) in

the valleys and hillsides.

The most common disturbance outside the National Park are farmers and urban uses while has an elevated recreational

pressure inside it.

.

Within the study area, according to the official classification provided by the National Park, there are 63 different plant

communities in accordance with their floristic composition. Given the variability of this classification, we decided

to group them to facilitate the statistical analysis of use versus availability (Table S1), setting out nine different

vegetation types (wet forest, thermophilic forest, rain forest, wooded areas, wet shrub, thermophilic shrub, wet

grassland, thermophilic grassland, and rocky areas with vegetation) (Table 1). 114

115

Capra pyrenaica 116

The Iberian ibex (Capra pyrenaica (Schinz, 1838) is an endemic wild ungulate of the Iberian Peninsula. It is distributed 117

in the major mountain ranges of the eastern and southern regions of the peninsula, as well as in Macizo de Gredos 118

(Herrero and Pérez 2008), thanks to numerous reintroductions during the second half of the 20th century (Refoyo et al. 119

2014). 120

121

In 1990 the reintroduction of the Iberian ibex began in the National Park of Sierra de Guadarrama with specimens of the 122

Capra pyrenaica victoriae (Cabrera 1911) from a preserve adjacent to the National Hunting Reserve of Gredos and Las 123

Batuecas National Hunting Reserve. A total of 67 ibex specimens (41 females and 26 males) were reintroduced in the 124

National Park until 1992 (Refoyo 2014). 125

126

Abundance and Density monitoring 127

Since reintroduction, the population was monitored during six field seasons (2000, 2003, 2005, 2007, 2010, and 2014) 128

by direct observation of the animals along transects using the distance sampling method (Buckland et al. 1993) This 129

method is useful when using the projected distance (Buckland et al. 2001), although some authors believe that the 130

transect method is not suitable to perform this kind of estimate for ungulates in very hilly areas (Carloti et al. 2015) due 131

to the difficulty of estimating the perpendicular distance value. We also calculated kilometric abundance indices (KAIs) 132

to abundance. .KAIs expresses the ratio of the total number of individuals observed along a transect by the total 133

transect length covered at each site. 134

135

For each contact, we recorded (using 8x40 to 10x50 binoculars) the number of individuals, habitat, sex, age of the 136

individuals, and the perpendicular distance to the transect line using a laser distance meter (Bushnell Yardage Pro 137

Sport). The study area progressively increased as the presence of the species was detected in new areas by forest 138

rangers (Forest rangers carry out regular monitoring of the presence of the species throughout the National Park 139

boundaries through monthly transects on foot and by car, and by the establishment of observation points. They note all 140

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the contacts detected of the species, including the detection of fecal samples); it occupied 4 590 ha during the field 141

seasons in 2000, 2003, 2005 and 2007, 5,764 ha during the 2010 field season, and 7 770 ha during the field season in 142

2014. The number of monitored kilometers varied each year with the increase of the surface (79 km in 2000, 2003, 143

2005, and 2007; 89 km in 2010; and 97 km in 2014). There were slight differences among the field seasons due to the 144

inaccessibility of certain areas and the difficulty of retrieving each of the preset transects. The same eight researchers 145

participated in all seasons. In 2000, 2003, 2005, and 2007 there were 22 transects, which had an average length of 3.64 146

km; in 2010 there were 2 more transects (24), which had an average length of 3.80 km; and in 2014 there were 5 more 147

transects (29), which had an average length of 3.29 km. There were slight differences among the field seasons due to 148

the inaccessibility of certain areas and the difficulty of retrieving each of the preset transects (Figure 1). All transects 149

were sampled on successive and climatically suitable days, either in the morning (2–3 hours after sunrise) or afternoon 150

(2–3 hours before sunset). The Distance 6.0 programme was used to calculate the population density (Thomas et al. 151

2009). This software was specifically designed to obtain animal population densities by the linear transect method 152

through observations from fixed points (Burnham et al. 1980). To adjust contacts to better distribution functions, 153

truncations distance (maximum detection distance of observation) was changed from 160 m to 350 m. 154

To determine the use that the species has on the different areas of the park throughout the year, the different field 155

seasons established for monitoring the population were adjusted to seasonal conditions, i.e. spring (June 2000, 2003, 156

and 2007), autumn (October 2005 and 2010), and summer (August 2014). We did not select any winter census data 157

since much of the territory was covered with snow and the availability of different habitats was reduced, forcing 158

population migration, which has already been detected in other wild ungulates (Garrott et al. 2003; Jacobson et al. 2004; 159

Wang et al. 2006). 160

161

Occupied area 162

Each of the obtained contacts in every field season was georeferenced and digitalised for subsequent treatment with 163

ArcMap 10.2 (Environmental Systems Research Institute, Inc. (Esri)) . We used a GPS for contact georeferencing (it 164

marks a point on the observer's position, i.e. a "waypoint"), scoring the distance from the contact to the observer (using 165

laser distance meters) and its relative orientation to geographic north. Later, and using Geographic Information Systems 166

(GIS) tools, we were able to relocate the obtained waypoints in the places where the contacts were detected. 167

168

To set the used and dispersal area, every contact was used considering the group size obtained during each field 169

season as an individual record of the population, indicative of the location preferences of the specimens. To establish 170

the degree of the population expansion, the distance of each contact was calculated in every campaign with respect to 171

the release point. To determine the occupied area, a kernel distribution analysis was performed (Silverman 1986; 172

Worton 1989). The occupied area for each field season included 50%, 75%, and 95% of the population. Because, the 173

occupation of space has relationship with the breeding habits of the species (in autumn the species conforms larger 174

groups so the occupied area tends to be lower, while in summer and spring has a greater occupation of the territory 175

since the groups were disgregated after the reproductive period) we consider only the no reproductive period (spring 176

and summer) . 177

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Kernel density estimation is a popular method for using a sample of points to estimate the distribution that generated 179

those points (Fortmann-Roe et al. 2012). This estimation technique is employed in ecology (Millspaugh et al. 2006) to 180

measure the intensity or probability of use throughout an animal's species distribution (Burt 1940), and also to measure 181

joint space use of multiple animals (Fieberg and Kochanny 2005; Daermon 2012). This technique is based on locations 182

obtained by telemetry (Gitzen et al. 2006). 183

184

Woody plants are known to be highly sensitive to herbivory because browsers can limit their regeneration (Perea and 185

Gil 2015). To determine the effects of ibex populations on vegetation in each season, an intersection was performed 186

between the vegetation layer and the kernel generated using the ArcMap 10.2 intersection tool (Environmental Systems 187

Research Institute, Inc. (Esri)). By this tool, it can be possible to know the total ha of each habitat present in the animal's 188

species distribution and its use over the years. 189

190

Statistical analysis 191

Different statistical analyses were performed using the General Regression Module of Statistica 7.0 (StatSoft Inc., 192

Tulsa, Oklahoma). To determine the expansion of the species, we performed a regression analysis considering the 193

distances of each contact to the release point in every field season. In order to identify the expansion of the species and 194

the differences between years, we conducted post-hoc tests. We selected Fisher's Least Significant Difference (LSD) 195

test. To determine differences in the use of the environment and availability, non-parametric analysis (chi-squared) was 196

performed. 197

RESULTS 198

The distance sampling models (key function and series expansion) that were selected for density estimations for the 199

three analyses were the same (semi-normal), but the statistical results were different. Truncating distance and interval 200

numbers are shown in Table 2. The density obtained for each field season showed a significant increase from 6.57 201

ind./km2 in 2000 (Refoyo 2014) to 44.82 ind./km2 in the last field season (2014) (Pablo Refoyo, Complutense University 202

of Madrid). Between the reintroduction and 2000, we noticed a population increase of 23%, followed by a 36% increase 203

between 2000 and 2003. Finally, the population increased by 19% annually from 2003 to 2007 (Refoyo et al. 2014); 204

between 2007 and 2014, the population increased by 9% annually (Table 2). 205

206

The area occupied by the species has been increasing over the past 15 years. If we consider the more intensively 207

occupied area (50% of the population), it has produced an increase of 150% from 715 ha in 2000 to 2130 ha in 2014. 208

This increase is greater (300%) if we consider 75% of the area occupied by the population, but only an increase of 209

100% when the occupation is 95% of the population (Table 3). 210

211

A linear relationship was detected between the abundance found in each field seasons and the extent of the area 212

occupied by the species within the National Park (Figure 2), especially with regard to the core area with the greater 213

presence of the species (area with 50% of contacts) (F: 90.059; p = 0.011). Density scores explain 97% of the variance 214

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of the occupied area during these years. This relationship holds when we consider 75% of the population (F: 26.675; p = 215

0.04) and explains 93% of the variance of the occupied area. This does not occur when considering 95% of the 216

population (F: 7.030; p = 0.12) (Figure 3). 217

218

The expansion of the species has occurred especially since 2010, i.e. during the last two field seasons (Figure 4). The 219

average distance to the release point at which the specimens were observed during each field season showed a 220

significant relationship (F: 10.845, df: 5: p< 0.001) that was greater than that from the 2010 field seasons (Scheffé test: 221

MS = 3003E3, df = 557; p < 0.001), i.e. when the population showed a higher density than 42 ind./km2 (Table 3). 222

223

During the spring, the area occupied by 50% of the population was 907 ha, which is about 11% of the studied area, 224

indicating a high concentration of the species. The vegetation in this area is broom with a surface of different degrees of 225

outcropping rock (61.5%), followed by pine (14%) and rocky areas (11%). In summer, the area occupied with a higher 226

concentration of specimens increased considerably and extended to 1 926 ha, more than double the species' 227

occupation in spring. In summer, it was found that the species mainly occupies the highlands of the study area. The 228

predominant vegetation in the area of greatest use is broom with rocky outcrops (71%), followed by pine forests (16%). 229

230

During the autumn, the occupied surface with a higher concentration of specimens was reduced by 20% compared to 231

the spring and stood at 764 ha. This decrease coincided with the behaviour of the species, as it tends to concentrate for 232

the breeding season (autumn–winter) and at lower altitudes than in spring, possibly influenced by climatic factors. The 233

vegetation of this area was similar to that in the spring, although the proportion of broom was reduced (44%) and the 234

use of pine (19%) and rocky areas (20%) was increased; the proportion of pasture land was maintained with respect to 235

the spring. In summary, the species is concentrated in less extensive areas in autumn, so pressure on it increases, and 236

is more extensive during the summer. The vegetation types that suffer more pressure from the species are broom, 237

followed by pine forests (Table 4). 238

239

When analysing the availability in relation to the use with respect to the season, we can see that the differences 240

between availability and use are significant for all seasons; however, while for spring (chi-squared = 47.17288; df = 7; p 241

<0.000) and summer (chi-squared = 74.74612; df = 7, p <0.000) the significance was high, in the case of autumn it was 242

not (chi-squared = 17.30653; df = 7; p <0.015525). In spring, scrub and wet grassland were more used than available, 243

while the thermophilic scrub, woodland areas, and thermophilic forests were infra-used. During summer, scrub areas 244

and wet grassland were used much more than available, while the scrublands, rocky areas, and thermophilic forest as 245

well as woodland areas were less used than was expected. In autumn, the differences between the observed and 246

expected were reduced, although bushes and rainforests were more used than expected (Table 4). 247

DISCUSSION 248

Values obtained in this work exceed the densities obtained for other populations of the genus Capra (Refoyo et al. 249

2014). Although due to the lack of data we do not know if these values are higher or not comparing with the data 250

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before the extinction. This population increase was higher than in other populations of the species as well as (Perez et 251

al., 2002) in other populations of genus Capra (Dupré et al. 2001, Carnevali et al. 2009) with increases about 3-6%, or in 252

others ungulates as roe deer (Capreolus capreolus) with increases about 4% (Calenge et al., 2005); possibly because 253

the species was reintroduced in a protected area without natural predators, with temperate climatic conditions (Toïgo et 254

al. 1997; Scillitani 2011) and without domestic flocks. 255

256

This increase is related to the increase in the area occupied by the species, initially only in relation to the main core 257

(core area), but later in a general way in recent years. As suggested by Vander Wal et al. (2014), saturation of 258

encounters and avoidance suggests that the benefits of social behaviour may become costs when two individuals share 259

a large extent of their home range. Besides, individuals compete over resources within a shared space (Rieucau and 260

Giraldeau 2011). On the other hand, a higher density could cause an increase in aggressive interactions as occurs with 261

other gregarious ungulates (Weckerly 1999). These factors could force the dispersion of some specimens and enhance 262

the rate of colonisation of new territories (Fuller 2007), as has happened with other ungulate populations (Messier et al. 263

1988). 264

265

Our results suggest that the population is properly settled in the area of reintroduction, as has also been found by other 266

authors (Messier et al. 1988), at least until they reach excessive densities. In this situation, there is a low dispersion of 267

the specimens that results in an excessive concentration in very specific areas, which is a factor that negatively affects 268

the vegetation (Refoyo et al. 2014) This low dispersion matches with the indicated by Pedrotti (1995) for the Alpine ibex 269

in the Alpi Orobie or for other ungulates as the roe deer (Capreolus capreolus) in mediterranean areas (Rossell et al., 270

1996; Calenge et al., 2005). 271

272

The benefits of social behaviour may become costs (Vander Wal et al. 2014) when the densities reach very high values 273

(44 ind./km2). For populations of group-living species, social structure likely has important demographic consequences 274

and can lead to dynamics that are qualitatively different from those of homogeneous populations (Bateman et al. 2012) 275

and that could force the species to faster dispersion. This pattern of increase to high density followed by expansion into 276

a new range is similar to that described for other ungulates (Larter et al. 2000). 277

278

It was found that the differential use of the different habitats present in the study area occurred depending on the time of 279

year, coinciding with what has been found for the species in the populations located in the Sierra de Gredos and Sierra 280

Nevada, where grasses are eaten more regularly in spring and early summer, but as the year progresses, woody 281

species gain prominence (Martinez 2002). Something similar happens in Puertos de Tortosa-Beceite, where woody 282

plants, in all seasons, are consumed in greater quantities than herbaceous plants, with emphasis in winter (88% of the 283

diet). Herbaceous plants are consumed more in spring and early summer (Martínez 1994). Petrotti (1995) indicates that 284

winter ranges characterize ibex as an insular species, especially by females and tend to remain in their traditional 285

ranges while increasing their density (Gauthier et al. 1994). 286

287

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Overall, in areas with abundant grasses (Gredos and Sierra Nevada) the species feeds on them, leaving woody plants 288

for winter when snow forces animals to a lower altitude. However, in areas with abundant woody plants (oak, pine, etc.) 289

the species feeds mainly on them (Cazorla, Beceite, and Tortosa), leaving grasses for times where these are abundant 290

(spring). 291

292

Differences detected between the availability and use of different habitats by the species may be due to the availability 293

of different trophic resources. As has been suggested in many studies, weather conditions are factors that affect the 294

processes of seasonal migration of many wild ungulates, mainly due to trophic resources (Gaillard et al. 2000; Clutton-295

Brock and Coulson 2002; Garrott et al. 2003; Jacobson et al. 2004; Wang et al. 2006). Range expansion may delay 296

responses to food limitation, such as diminished survival and fecundity, until new areas can no longer be colonised to 297

provide additional forage (Messier et al. 1988; Larter et al. 2000), forcing the population to migrate (Fuller et al. 2007). 298

Ungulate populations generally become more sensitive to density-independent factors that affect resource availability as 299

they approach high densities (Saether 1997; Gaillard et al. 1998, 2000). 300

301

Finally, the high density of ungulates can cause excessive pressure on vegetation (Palacios et al. 1989; Álvarez and 302

Ramos 1991; Perea et al. 2014). High densities of ibex in a protected area may threaten certain endangered plant 303

species (Perea et al. 2015), so continuous monitoring of the population is a very effective tool to know the evolution of it 304

and establish the control measures that are necessary to reduce the population to environmentally acceptable levels. 305

306

The Iberian ibex population increase has been the cause of changes in the fauna and flora of the National Park of Sierra 307

de Guadarrama. Our data allow us to demonstrate the importance of monitoring reintroduced populations and 308

establishing the need to control the reintroduced populations to avoid these alterations in a protected area. It is 309

necessary to properly manage reintroduced populations in order to allow the coexistence of the different species in 310

protected areas. 311

Protected areas are essential for the reintroductions of Iberian ibex because they improve the probability of success 312

(Pérez et al, 2002; Moço et al., 2006; Carnevali et al., 2009; Goldstein and Rominger 2011; Refoyo et al., 2012), 313

however, as it is demonstrated in this work, the new reintroductions should consider the low dispersion of the specimens 314

as their seasonal distribution to reduce effects on threatened plant species. It is also essential to know the capacity of 315

the environment and the establishment of control measures to prevent the population to acquire densities that the 316

environment can not support. 317

Acknowledgments 318

This study was possible thanks to the collaboration of those in charge of the former Regional Park of the Cuenca Alta 319

del Manzanares, now the National Park of the Sierra de Guadarrama, and the technical staff of the company Estudios 320

Territoriales Integrados, S.L., who collaborated with us in data collection. 321

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493

494

495

496

Table 1: Available area in ha and % of total surface of the National Park for each

established habitat and Total area. We group the habitat into nine different vegetation

types (Table S1) (wet forest, thermophilic forest, rain forest, wooded areas, wet shrub,

thermophilic shrubs, wet grassland, thermophilic grassland, and rocky areas with

vegetation). 497

Habitat Available area

Ha %

Wet forest 14.2 0.18

Thermophilic forest 873.1 11.24

Rain forest 1099.6 14.15

Wooded area 767.7 9.88

Thermophilic scrub 496.9 6.39

Wet brushwood 2682.7 34.53

Rocky area 1453.4 18.70

Wet grassland 312.5 4.02

Xerophilous grassland 55.5 0.71

Urban Areas 14.6 0.19

Total 7770.2 100

498

499

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Table 2. Evolution of the number of animals and density of Iberian ibex (Capra 500

pyrenaica, Schinz, 1838) between 2000 and 2014 in Sierra de Guadarrama National 501

Park. KIAs. Abundace index; f(0) value of probability density function at zero for line 502

transects = 1/u (u = W*p; W = width of line transect; p = probability of observing an 503

object in defined area. Truncating distance (m) = maximum detection distance. 504

Year Number

of

animals

KIAs

(ind/km)

Density

(ind/km2)

Coefficient

of variation

(CV%)

Truncating

distance (m)

Interval

numbers

f(0) p Degrees

of

freedom

95% confidence

interval

2000 359 4.16 6.67 38.47 350 3 0.0098 0.29 39 3.149 14.146

2003 773 5.69 16.83 25.50 300 3 0.0090 0.36 117 10.238 27.669

2005 1065 6.06 23.2 25.70 300 4 0.011 0.30 62 13.992 38.447

2007 1523 5.74 33.16 25.06 160 6 0.014 0.44 32 20.129 55.016

2010 2437 11.09 42.29 20.62 240 6 0.013 0.40 198 28.190 63.430

2014 3324 8.65 42.88 20.81 200 4 0.012 0.39 216 28.5 64.3

505

506

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Table 3: Occupied area (ha) by 50%, 75% and 95% of the species distribution using the 507

Kernel distribution by years (in parenthesis monitoring season). 508

509

Field seasons Ocuppied area (ha)

50% 75% 95%

2000 (spring) 715 1,375 4,590

2003 (spring) 1,004 1,866 4,590

2005 (autumn) 1,101 1,956 4,590

2007 (spring) 1,169 2,255 4,590

2010 (autumn) 1,609 3,255 5,764

2014 (summer) 2,130 5,472 7,770

510

511

512

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513

Table 4: Total available area by Habitat present in the National Park and available area 514

by Habitat used by ibex (Capra pyrenaica, Schinz, 1838) populations in each seasons 515

(in ha and %). 516

Habitat Available area

Habitat used

Spring

Habitat used

Summer

Habitat used

Autumn

Ha % Ha % ha % ha %

Wet forest 14.2 0.18 0 0 0 0 0 0

Thermophilic

forest 873.1 11.24 46.77 5.16 4.00 0.21 56.87 7.45

Rain forest 1099.6 14.15 124.90 13.78 301.09 15.82 147.94 19.38

Wooded area 767.7 9.88 0.36 0.04 69.47 3.65 43.82 5.74

Thermophilic

scrub 496.9 6.39 26.10 2.88 13.51 0.71 28.93 3.79

Wet brushwood 2682.7 34.53 557.42 61.5 1345.60 70.7 338.70 44.37

Rocky area 1453.4 18.70 75.68 8.35 39.02 2.05 84.05 11.01

Wet grassland 312.5 4.02 75.14 8.29 119.33 6.27 63.05 8.26

Xerophilous

grassland 55.5 0.71 0 0 0 0 0 0

Urban Areas 14.6 0.19 0 0 0 0 0 0

Total 7770.2 100 906.38 100 1903.25 99.41 763.36 100

517

518

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Figure 1: Study area with the boundaries of the National Park of Sierra de Guadarrama; 519

Samples area, Transects and Reintroduction point. 520

Figure 2: Occupied area for each field seasons included 50%, 75% and 95% of the 521

population by Kernel distribution The occupied area is represented in gradient gray with 522

darker maximal values (95%). and clearer lower (50%). It is also indicated the 523

reintroduction point. 524

Figure 3: Relationship between abundance index (KAI,s) in no reproductive field 525

seasons and occupied area by 50, 75 and 95% of the population by Kernel distribution. 526

Figure 4: Relationship between Field seasons and distance form releasing point found in 527

each field seasons and the extent of the area occupied by the species within the National 528

Park (post-hoc Scheffé test: MS = 3003E3, df = 557; p < 0.001 (Distance/field seasons) 529

530

531

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Figure 1: Study area with the boundaries of the National Park of Sierra de Guadarrama; 532

Samples area, Transects and Reintroduction point. 533

534

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Figure 2: Occupied area for each field seasons included 50%, 75% and 95% of the 535

population by Kernel distribution The occupied area is represented in gradient gray with 536

darker maximal values (95%). and clearer lower (50%). It is also indicated the 537

reintroduction point. 538

539

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540

Figure 3: Relationship between abundance index (KAI,s) in no reproductive field 541

seasons and occupied area by 50, 75 and 95% of the population by Kernel distribution. 542

543

544

50% 75% 95%3 4 5 6 7 8 9

KAI,s

Field seasons

2000 2003 2007 2014

0

2000

4000

6000

8000

10000

12000

14000

Occupied area (ha)

Ika;(core area 50%): r2 = 0,9783; r = 0,9891; p = 0,0109

Ika; (core area75%): r2 = 0,9303; r = 0,9645; p = 0,0355

Ika; (core area 95%): r2 = 0,7785; r = 0,8823; p = 0,1177

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545

Figure 4: Relationship between Field seasons and distance form releasing point found in 546

each field seasons and the extent of the area occupied by the species within the National 547

Park (post-hoc Scheffé test: MS = 3003E3, df = 557; p < 0.001 (Distance/field seasons) 548

549

Current effect: F(5, 557)=10,845, p=,00000Vertical bars denote 0,95 confidence intervals

2000 2003 2005 2007 2010 2014

Field seasons

2000

2500

3000

3500

4000

4500

5000

DISTANCE (m.)

Page 25 of 25



Table S1. Habitat grouped into nine different vegetation types (wet forest, thermophilic forest, rain forest, wooded

areas, wet shrub, thermophilic shrub, wet grassland, thermophilic grassland, and rocky areas with vegetation).

Habitat ha

Wet

fore

st TOTAL 14.2

Pyrenean oak trees with <5% of rock outcropping with a cover of 25%–50% with grassland 13.2

Pyrenean oak trees with 5%–25% of rock outcropping with a 25%–50% covered with grassland 1

Th

erm

ophili

c f

ore

st

TOTAL 873.1

Mixed pine forests of an adult repopulation of Pinus pinaster and P. nigra with 25%–50% of rock outcropping; >50% pine

cover with brushwood 195.4

Mixed pine forests of Pinus pinaster and an adult repopulation of P. nigra with 5%–25% of rock outcropping; 20%–50%

pine cover with brushwood 5.2

Mixed natural pine forests of Pinus pinaster, P. sylvestris and P.nigra with 5%–25% of rock outcropping; >50% pine cover

with brushwood 2.7

Mixed pine forest of Pinus pinaster, P. sylvestris and a repopulation of P. nigra with 25%–50% of rock outcropping; >50%

repopulation pine's cover with brushwood and grassland 284.6

Mixed pine forests of Pinus sylvestris and natural P. nigra with 25%–50% of rock outcropping; >50% pine cover with 20%–

50% trees 291.6

Mixed pine forests of Pinus sylvestris and an adult repopulation of P. nigra with 25%–50% of rock outcropping; >50% pine

cover with brushwood 80.7

Natural pine forests of Pinus pinaster with 25%–50% of rock outcropping; 20%–50% pine cover with brushwood and rock

rose 5.6

Pine forest of a heterogeneus repopulation of Pinus pinaster with 25%–50% of rock outcropping; 20%–50% pine cover with

brushwood and rock rose 7.3

Rain

fore

st

TOTAL 1099.6

Natural pine forests of Pinus sylvestris with 25%–50% of rock outcropping; >50% pine cover with brushwood and broom 694.6

Natural pine forests of Pinus sylvestris with 5%–25% of rock outcropping; >50% pine cover with brushwood 128.4

Natural pine forests of Pinus sylvestris with 25%–50% of rock outcropping; >50% pine cover with brushwood and ferns 41.8

Pine forests of an adult repopulation of Pinus sylvestris with 5%–25% of rock outcropping; >50% pine cover with

brushwood 11.4

Pine forests of a repopulation of Pinus sylvestris with <5% of rock outcropping; >50% repopulation cover with

brushwood,broom and Pinus uncinata 1.6

Pine forests of a repopulation of Pinus sylvestris with <5% of rock outcropping; >50% pure with broom, pasture and Pinus

uncinata 0.2

Pine forests of a repopulation of Pinus sylvestris with 25%–50% of rock outcropping; >50% repopulation cover with >50%

of grassland 21

Pine forests of a repopulation of Pinus sylvestris with rock outcropping; 25%–50% of failed repopulations with brushwood,

broom and grassland 196.8

Pine forests of a repopulation of Pinus sylvestris with rock outcropping; 5%–25% of repopulation cover with brushwood 3.8

Wooded a

reas

TOTAL 767.7

Rocky area with cover grade >50% with trees, 5%–20% brushwood with holm oak 184.2

Rocky area with cover grade >50% with trees, 5%–20% brushwood with holm oak and rockrose 43.9

Rocky area with cover grade >50% with trees, 5%–20% brushwood with juniper 273.2

Rocky area with cover grade >50% with trees, 5%–20% brushwood with Pinus sylvestris 88

Rocky area with cover grade >50% with trees, 5%–20% brushwood with Pinus sylvestris and Pinus uncinata. 178.4

Th

erm

ophili

c s

cru

b

TOTAL 1950.3

Lavenders, thyme and other acidophilus plants of small size with rock outcropping; 5%–25% pure with patches of grass,

with Pyrenean oak and ash 26.1

Juniper with 25%–50% rock outcropping with 25%–50% cover with brushwood, lavenders and ferns. 28.6

Rockroses with rock outcropping; 25%–50% pure with patches of grassland 131.7

Rockroses with rock outcropping; 25%–50% pure with patches of pastures and junipers. 295.5

Rockroses with rock outcropping; 5%–25% mosaic with trees and/or shrub species (>50%) with patches of wasteland, with

oaks and ferns

15

Rocky area with cover grade >50% without wooded thicket, with brushwood and rockroses 217.7

Rocky area with cover grade >50% without wooded thicket, with rockroses and junipers 858.4

Rocky area with cover grade >50% without wooded thicket, with brooms 332.2

Rocky area with cover grade >50% without wooded thicket, with brooms and bearberry 45.1

Wet

bru

shw

ood

TOTAL 2682.7

Acidophilous mountainous brushwood with predominance of legumes with rock outcropping; 25%–50% mosaic of trees

and shrub species (>50% brushwood) with patches of grassland and Pinus sylvestris 7

Brooms and other bushes with rock outcropping; <5% of grassland 52

Brooms and other bushes with rock outcropping; <5% pure 14.9

Brooms and other bushes with rock outcropping; 25%–50% with grassland 565.2

Brooms and other bushes with rock outcropping; 25%–50% of grassland, with kermes oak and holm oak 116.7

Brooms and other bushes with rock outcropping with 25%–50% of grassland with juniper 275.7

Brooms and other bushes with rock outcropping with 25%–50% of grassland with Pinus sylvestris 80.7

Brooms and other bushes with rock outcropping; 25%–50% pure 566

Brooms and other bushes with rock outcropping; 5%–25% with grassland 282.2

Brooms and other bushes with rock outcropping; 5%–25% with grassland and ferns 484.4

Brooms and other bushes with rock outcropping; 5%–25% with grassland and with juniper 48

Brooms and other bushes with rock outcropping; 5%–25% pure 189.9

Wet

gra

ssla

nd

TOTAL 312.5

Cervum grazes (Nardus stricta) and wet grasslands with rock outcropping; 5%–25% pure with scrub and /or ferns and

brooms 103

Cervum grazes (Nardus stricta) and wet grasslands with rock outcropping; 5%–25% pure with with scrub and/or ferns 27.8

Cervum grazes (Nardus stricta) and wet grasslands with pure rock outcropping <5% 2.8

Brachypodium sp. with rock outcropping; <5% pure with scrubs and/or ferns with brooms 67.1

Brachypodium sp. with rock outcropping; 25%–50% pure with scrubs and/or ferns with brooms 50.6

Brachypodium sp. with rock outcropping; 5%–25% mixed of scrubs and/or ferns 15.3

Brachypodium sp. with rock outcropping; 5%–25% mixed of scrubs and/or ferns, with brooms 16.5

Brachypodium sp. with rock outcroppings; 5%–25% pure with scrubs and/or ferns 29.4

Xero

philo

us g

rassla

nd

TOTAL 55.5

Mesophyll grassland with rock outcropping; <5% pure with trees 4.1

Mesophyll grassland with rock outcropping; <5% pure with scrubs and/or ferns 6.8

Mesophyll grassland with rock outcropping; 5%–25% pure, with scrub and/or ferns and woodland 2.8

Xerophytic grassland with rock outcropping; <5% pure with brushwood and/or ferns and woodland 5.1

Xerophytic grassland with rock outcropping; 25%–50% pure with scrub and/or fern 32.5

Xerophytic grassland with rock outcropping; 5%–25% pure with lavenders and pure with juniper

4.2

Urb

an a

reas

TOTAL 14.6

Sports areas 5.2

Family houses and townhouses 5.8

Urban areas and urbanized areas 2.7

Other (camping, cemeteries,…) 0.9

OVERALL TOTAL 7770.2

draft - tspace repository: home · 2016-02-26 · draft 2 13 space use of a reintroduced population...

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