Canadian Journal of Zoology
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 AntropologyDraft Muñoz, Benito; Complutense University of Madrid, Zoology and Physical Antropology
Capra pyrenaica, Iberian Ibex, density-dependent, Reintroduction, National Keyword: Park, Spain
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1 Space use of a reintroduced population of Capra
2 pyrenaica in a protected natural area
3 P. Refoyo (Corresponding author); C. Olmedo and B. Muñoz.
4
5 P. Refoyo. Department of Zoology and Physical Anthropology, Complutense University of Madrid. C/ José Antonio
6 Novais, 12, E-28040 Madrid. PHONE: +34 91 394 50 31 FAX: 34 91 394 49 47 [email protected] (Corresponding
7 author)
8 C. Olmedo. Department of Zoology and Physical Anthropology, Complutense University of Madrid. C/ José Antonio
9 Novais, 12, E-28040 Madrid. [email protected]
10 B. Muñoz. Department of Zoology and Physical Anthropology, Complutense University of Madrid. C/ José Antonio
11 Novais, 12, E-28040 Madrid. [email protected]
12 Draft
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13 Space use of a reintroduced population of Capra pyrenaica in a protected natural area
14 P. Refoyo (Corresponding author); C. Olmedo and B. Muñoz.
15 ABSTRACT
16 In Europe, wild ungulates have undergone major expansion and population growth during recent decades. In certain
17 cases, the high density achieved by these populations has led to excessive pressure on the environment, which
18 eventually becomes a limiting factor for the population itself. One of these reintroductions was performed with the
19 Iberian ibex ( Capra pyrenaica (Schinz, 1838)) in the National Park of the Sierra de Guadarrama (Spain). This
20 reintroduced population was monitored during six field seasons (2000, 2003, 2005, 2007, 2010, and 2014) by direct
21 observation of the animals along transects using the distance sampling method to determine the degree of expansion
22 over the years and the use of different habitats according to different seasons. The abundances obtained for each field
23 season showed a significant increase from 4.16 ind./km to 8.65 ind./km, showing a linear relationship between the
24 abundance and the extent of the area occupied by the species. We observed that differences between habitat
25 availability and use were significant for all seasons.. Our data can be used as an example of the process of colonisation
26 of a population of wild ungulates and their impact on vegetation in order to better manage future reintroductions. 27 Draft 28 Keywords : Capra pyrenaica, Iberian Ibex, density-dependent, Reintroduction, National Park, Spain. 29
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30 INTRODUCTION
31 The introduction or reintroduction of ungulates is closely linked to human activities and movements (Christie and
32 Andrews 1966), Some of these reintroductions have been successful, as the case of the Oryx (Oryx sp.) In Oman or
33 ibex ((Capra ibex) in the Alps or Capra pyrenaica in Peneda-Gerês National Park in northern Portugal (Moço et al.,
34 2006), however others have failed as the attempts to introduce reindeer (Rangifer tarandus) in Matthews Island (Bering
35 Sea) (Soriguer et al., 1998). In all cases, these reintroductions should be done as suggested by international
36 organizations (IUCN 1998) that highlights the importance of theparameter of parameters of the population evolution
37 (Converse et al. 2013) and their relationship with the environment (Chapman 1928; Odum 1986).
38 This study goes into detail about the dispersion and use of space of a wild ungulate’s reintroduced population in a
39 Natural Protected Area.
40
41 In Europe, wild ungulates have undergone major expansion and population growth in recent decades, mainly going from
42 marginal areas to occupying a large part of the territory (Apollonio et al. 2010). There have been numerous
43 reintroductions of ungulates in protected natural areas (Geremia et al. 2011) that have enabled better protection of
44 these populations. 45 Draft 46 However, these protected natural spaces become restricted areas from where individuals cannot leave without
47 increasing the risk of their survival. This situation leads to a high concentration of ungulates in very limited areas and
48 causes high pressure on vegetation, which becomes a limiting factor affecting the dynamics of the population (Gaillard
49 et al. 2000; Bonenfant et al. 2009). On the other hand, a high density has significant costs for the population (Côté et al.
50 1995; Loehle 1995; Stillman et al. 1997; Krause et al. 2002; Fortin et al. 2009), and it can have important implications in
51 terms of social behaviour (Bateman et al. 2012), survival, and reproduction (Gaillard et al. 1998, 2000; Eberhardt 2002;
52 Festa-Bianchet et al. 2003).
53
54 In our case, the Iberian ibex (Capra pyrenaica (Schinz, 1838)) was distributed in the major mountain ranges of the
55 middle, eastern and southern of the Peninsula until the late nineteenth century. Excessive pressure exerted by men
56 produced its disappearance in much of the territory (Cabrera 1914; Perez et al. 2002).
57
58 The reintroduction of the Iberian ibex in Sierra de Guadarrama National Park began in 1990 with specimens of Capra
59 pyrenaica victoriae Cabrera, 1911 subspecies from a preserve adjacent to the National Hunting Reserve of Gredos and
60 Las Batuecas National Hunting Reserve. A total of 67 specimens (41 females and 26 males) were reintroduced by the
61 time it was finalized in 1992. The average age of the population was around 5 years, and the sex ratio of the introduced
62 specimens favored females at a proportion close to 1.6:1 (41 females and 26 males). However, among juveniles (under
63 5 years), the sex ratio favored males 1:1.4. On the contrary, there was a predominance of females in the individuals
64 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
65 other ibex reintroductions (Girard et al. 1998).
66
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67 Emigration and range expansion have been documented in several large ungulate populations when forage quantity or
68 quality has decreased due to density-dependent resource consumption (Lemke et al. 1998; Aanes et al. 2000; Larter et
69 al. 2000; Ferguson et al. 2001; Amarasekare 2004). There are also many studies of modern animal movement relative
70 to partial migration (Hebblewhite and Merrill 2009; Rivrud et al. 2010); however, most of these studies have focused on
71 natural populations, and few have bothered to analyse the process of land occupation of a reintroduced population, as
72 in our case.
73
74 The relationship between habitat selection and the abundance and distribution of species in a particular area has also
75 been widely discussed by several authors (Johnson 1980; Rosenzweig 1981) and constitutes an integral part of
76 effective management and conservation (Boyce and McDonald 1999). Habitat selection occurs over large areas when
77 we consider higher order scales of selection (Johnson 1980; Kittle et al. 2008; Peters et al. 2013), but also very small
78 spatial scales (e.g. warblers in a tree: MacArthur 1958). Similarly, habitat selection can vary over time (e.g. seasons)
79 (Singer 1979; Jenkins and Wright 1988; Macandza et al. 2012) as the forage base is progressively depleted (Brown and
80 Rosenzweig 1986; Van Beest et al. 2010). Nevertheless, there have not been many studies focusing on the effective
81 impact on the core area considering the seasonal movements of the species.
82 83 The aim of this work was to establish changes Draftof the occupied area of the reintroduced Iberian ibex during settlement 84 and colonisation. Our goals were: (1) to determine if there has been an increase in the area of distribution of the species
85 during the settlement process, (2) to determine whether there are relationships between the area of occupation and the
86 detected population increase during the settlement process, (3) to establish if the occupation of the area by ibex differs
87 seasonally and to determine if the pressure on the different plant communities in the park continues throughout the year,
88 and (4) to confirm if there is habitat selection regardless of availability in each season.
89
90 Our data can be used as an example of the process of colonisation of a population of wild ungulates.
91
92 MATERIALS AND METHODS
93 Study area
94 We monitored Iberian ibex in the National Park of the Sierra de Guadarrama. This National Park (Figure 1) has an area
95 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
96 shows a marked difference in altitude (between 2.200 and 1.100 m), alternating between very steep rocky areas (“Las
97 Pedrizas”) and areas of gentle topography. In general, they are very old rocks (from Paleozoic and Mesozoic) such
98 asgneisses, marbles and schists. Sierra de Guadarrama’s climate is typically continental with significant temperature
99 variations between seasons and very dry summers. The average temperature of the coldest month is -0.7°C and 16.4°C
100 in the warmest month, (January and July respectively). The annual rainfall is 1,325 mm with a high seasonal variation,
101 being July (24.9 mm) and November (176.5 mm) the driest months On average, the days of snowfall are 78 and 142 the
102 frost days a year.. The vegetation growing period is 5 months, although is less in the peaks above 2,000 meters .. Its
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103 vegetation includes shrubs (Cytisus purgans, Juniperus communis nana) and grassland (Festuca indigesta, Nardus 104 stricta, Festuca rubra) in highland areas, areas of Mediterranean shrubs (Cistus ladanifer, Rosmarinus officinalis, 105 Thymus vulgaris, Lavandula stoechas) in steeply sloped areas, and forests (Quercus ilex, Q. pyrenaica, Pinus spp.) in 106 the valleys and hillsides. 107 The most common disturbance outside the National Park are farmers and urban uses while has an elevated recreational 108 pressure inside it. 109 . 110 Within the study area, according to the official classification provided by the National Park, there are 63 different plant 111 communities in accordance with their floristic composition. Given the variability of this classification, we decided
112 to group them to facilitate the statistical analysis of use versus availability (Table S1), setting out nine different 113 vegetation types (wet forest, thermophilic forest, rain forest, wooded areas, wet shrub, thermophilic shrub, wet 114 grassland, thermophilic grassland, and rocky areas with vegetation) (Table 1). 115
116 Capra pyrenaica
117 The Iberian ibex (Capra pyrenaica (Schinz, 1838) is an endemic wild ungulate of the Iberian Peninsula. It is distributed
118 in the major mountain ranges of the eastern and southern regions of the peninsula, as well as in Macizo de Gredos 119 (Herrero and Pérez 2008), thanks to numerousDraft reintroductions during the second half of the 20th century (Refoyo et al. 120 2014). 121
122 In 1990 the reintroduction of the Iberian ibex began in the National Park of Sierra de Guadarrama with specimens of the
123 Capra pyrenaica victoriae (Cabrera 1911) from a preserve adjacent to the National Hunting Reserve of Gredos and Las
124 Batuecas National Hunting Reserve. A total of 67 ibex specimens (41 females and 26 males) were reintroduced in the
125 National Park until 1992 (Refoyo 2014). 126
127 Abundance and Density monitoring
128 Since reintroduction, the population was monitored during six field seasons (2000, 2003, 2005, 2007, 2010, and 2014)
129 by direct observation of the animals along transects using the distance sampling method (Buckland et al. 1993) This
130 method is useful when using the projected distance (Buckland et al. 2001), although some authors believe that the
131 transect method is not suitable to perform this kind of estimate for ungulates in very hilly areas (Carloti et al. 2015) due
132 to the difficulty of estimating the perpendicular distance value. We also calculated kilometric abundance indices (KAIs)
133 to abundance. .KAIs expresses the ratio of the total number of individuals observed along a transect by the total
134 transect length covered at each site. 135
136 For each contact, we recorded (using 8x40 to 10x50 binoculars) the number of individuals, habitat, sex, age of the
137 individuals, and the perpendicular distance to the transect line using a laser distance meter (Bushnell Yardage Pro
138 Sport). The study area progressively increased as the presence of the species was detected in new areas by forest
139 rangers (Forest rangers carry out regular monitoring of the presence of the species throughout the National Park
140 boundaries through monthly transects on foot and by car, and by the establishment of observation points. They note all
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141 the contacts detected of the species, including the detection of fecal samples); it occupied 4 590 ha during the field
142 seasons in 2000, 2003, 2005 and 2007, 5,764 ha during the 2010 field season, and 7 770 ha during the field season in
143 2014. The number of monitored kilometers varied each year with the increase of the surface (79 km in 2000, 2003,
144 2005, and 2007; 89 km in 2010; and 97 km in 2014). There were slight differences among the field seasons due to the
145 inaccessibility of certain areas and the difficulty of retrieving each of the preset transects. The same eight researchers
146 participated in all seasons. In 2000, 2003, 2005, and 2007 there were 22 transects, which had an average length of 3.64
147 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
148 transects (29), which had an average length of 3.29 km. There were slight differences among the field seasons due to
149 the inaccessibility of certain areas and the difficulty of retrieving each of the preset transects (Figure 1). All transects
150 were sampled on successive and climatically suitable days, either in the morning (2–3 hours after sunrise) or afternoon
151 (2–3 hours before sunset). The Distance 6.0 programme was used to calculate the population density (Thomas et al.
152 2009). This software was specifically designed to obtain animal population densities by the linear transect method
153 through observations from fixed points (Burnham et al. 1980). To adjust contacts to better distribution functions,
154 truncations distance (maximum detection distance of observation) was changed from 160 m to 350 m.
155 To determine the use that the species has on the different areas of the park throughout the year, the different field
156 seasons established for monitoring the population were adjusted to seasonal conditions, i.e. spring (June 2000, 2003, 157 and 2007), autumn (October 2005 and 2010), andDraft summer (August 2014). We did not select any winter census data 158 since much of the territory was covered with snow and the availability of different habitats was reduced, forcing
159 population migration, which has already been detected in other wild ungulates (Garrott et al. 2003; Jacobson et al. 2004;
160 Wang et al. 2006).
161
162 Occupied area
163 Each of the obtained contacts in every field season was georeferenced and digitalised for subsequent treatment with
164 ArcMap 10.2 (Environmental Systems Research Institute, Inc. (Esri)) . We used a GPS for contact georeferencing (it
165 marks a point on the observer's position, i.e. a "waypoint"), scoring the distance from the contact to the observer (using
166 laser distance meters) and its relative orientation to geographic north. Later, and using Geographic Information Systems
167 (GIS) tools, we were able to relocate the obtained waypoints in the places where the contacts were detected.
168
169 To set the used and dispersal area, every contact was used considering the group size obtained during each field
170 season as an individual record of the population, indicative of the location preferences of the specimens. To establish
171 the degree of the population expansion, the distance of each contact was calculated in every campaign with respect to
172 the release point. To determine the occupied area, a kernel distribution analysis was performed (Silverman 1986;
173 Worton 1989). The occupied area for each field season included 50%, 75%, and 95% of the population. Because, the
174 occupation of space has relationship with the breeding habits of the species (in autumn the species conforms larger
175 groups so the occupied area tends to be lower, while in summer and spring has a greater occupation of the territory
176 since the groups were disgregated after the reproductive period) we consider only the no reproductive period (spring
177 and summer) .
178
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179 Kernel density estimation is a popular method for using a sample of points to estimate the distribution that generated
180 those points (Fortmann-Roe et al. 2012). This estimation technique is employed in ecology (Millspaugh et al. 2006) to
181 measure the intensity or probability of use throughout an animal's species distribution (Burt 1940), and also to measure
182 joint space use of multiple animals (Fieberg and Kochanny 2005; Daermon 2012). This technique is based on locations
183 obtained by telemetry (Gitzen et al. 2006).
184
185 Woody plants are known to be highly sensitive to herbivory because browsers can limit their regeneration (Perea and
186 Gil 2015). To determine the effects of ibex populations on vegetation in each season, an intersection was performed
187 between the vegetation layer and the kernel generated using the ArcMap 10.2 intersection tool (Environmental Systems
188 Research Institute, Inc. (Esri)). By this tool, it can be possible to know the total ha of each habitat present in the animal's
189 species distribution and its use over the years.
190
191 Statistical analysis
192 Different statistical analyses were performed using the General Regression Module of Statistica 7.0 (StatSoft Inc.,
193 Tulsa, Oklahoma). To determine the expansion of the species, we performed a regression analysis considering the
194 distances of each contact to the release point in every field season. In order to identify the expansion of the species and 195 the differences between years, we conducted post-hocDraft tests. We selected Fisher's Least Significant Difference (LSD) 196 test . To determine differences in the use of the environment and availability, non-parametric analysis (chi-squared) was
197 performed.
198 RESULTS
199 The distance sampling models (key function and series expansion) that were selected for density estimations for the
200 three analyses were the same (semi-normal), but the statistical results were different. Truncating distance and interval
201 numbers are shown in Table 2. The density obtained for each field season showed a significant increase from 6.57
202 ind./km 2 in 2000 (Refoyo 2014) to 44.82 ind./km 2 in the last field season (2014) (Pablo Refoyo, Complutense University
203 of Madrid). Between the reintroduction and 2000, we noticed a population increase of 23%, followed by a 36% increase
204 between 2000 and 2003. Finally, the population increased by 19% annually from 2003 to 2007 (Refoyo et al. 2014);
205 between 2007 and 2014, the population increased by 9% annually (Table 2).
206
207 The area occupied by the species has been increasing over the past 15 years. If we consider the more intensively
208 occupied area (50% of the population), it has produced an increase of 150% from 715 ha in 2000 to 2130 ha in 2014.
209 This increase is greater (300%) if we consider 75% of the area occupied by the population, but only an increase of
210 100% when the occupation is 95% of the population (Table 3).
211
212 A linear relationship was detected between the abundance found in each field seasons and the extent of the area
213 occupied by the species within the National Park (Figure 2), especially with regard to the core area with the greater
214 presence of the species (area with 50% of contacts) ( F: 90.059; p = 0.011). Density scores explain 97% of the variance
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215 of the occupied area during these years. This relationship holds when we consider 75% of the population ( F: 26.675; p =
216 0.04) and explains 93% of the variance of the occupied area. This does not occur when considering 95% of the
217 population ( F: 7.030; p = 0.12) (Figure 3).
218
219 The expansion of the species has occurred especially since 2010, i.e. during the last two field seasons (Figure 4). The
220 average distance to the release point at which the specimens were observed during each field season showed a
221 significant relationship ( F: 10.845, df: 5: p< 0.001) that was greater than that from the 2010 field seasons (Scheffé test:
222 MS = 3003E3, df = 557; p < 0.001), i.e. when the population showed a higher density than 42 ind./km 2 (Table 3).
223
224 During the spring, the area occupied by 50% of the population was 907 ha, which is about 11% of the studied area,
225 indicating a high concentration of the species. The vegetation in this area is broom with a surface of different degrees of
226 outcropping rock (61.5%), followed by pine (14%) and rocky areas (11%). In summer, the area occupied with a higher
227 concentration of specimens increased considerably and extended to 1 926 ha, more than double the species'
228 occupation in spring. In summer, it was found that the species mainly occupies the highlands of the study area. The
229 predominant vegetation in the area of greatest use is broom with rocky outcrops (71%), followed by pine forests (16%).
230 231 During the autumn, the occupied surface with aDraft higher concentration of specimens was reduced by 20% compared to 232 the spring and stood at 764 ha. This decrease coincided with the behaviour of the species, as it tends to concentrate for
233 the breeding season (autumn–winter) and at lower altitudes than in spring, possibly influenced by climatic factors. The
234 vegetation of this area was similar to that in the spring, although the proportion of broom was reduced (44%) and the
235 use of pine (19%) and rocky areas (20%) was increased; the proportion of pasture land was maintained with respect to
236 the spring. In summary, the species is concentrated in less extensive areas in autumn, so pressure on it increases, and
237 is more extensive during the summer. The vegetation types that suffer more pressure from the species are broom,
238 followed by pine forests (Table 4).
239
240 When analysing the availability in relation to the use with respect to the season, we can see that the differences
241 between availability and use are significant for all seasons; however, while for spring (chi-squared = 47.17288; df = 7; p
242 <0.000) and summer (chi-squared = 74.74612; df = 7, p <0.000) the significance was high, in the case of autumn it was
243 not (chi-squared = 17.30653; df = 7; p <0.015525). In spring, scrub and wet grassland were more used than available,
244 while the thermophilic scrub, woodland areas, and thermophilic forests were infra-used. During summer, scrub areas
245 and wet grassland were used much more than available, while the scrublands, rocky areas, and thermophilic forest as
246 well as woodland areas were less used than was expected. In autumn, the differences between the observed and
247 expected were reduced, although bushes and rainforests were more used than expected (Table 4).
248 DISCUSSION
249 Values obtained in this work exceed the densities obtained for other populations of the genus Capra (Refoyo et al.
250 2014). Although due to the lack of data we do not know if these values are higher or not comparing with the data
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251 before the extinction. This population increase was higher than in other populations of the species as well as (Perez et
252 al., 2002) in other populations of genus Capra (Dupré et al. 2001, Carnevali et al. 2009) with increases about 3-6%, or in
253 others ungulates as roe deer ( Capreolus capreolus ) with increases about 4% (Calenge et al., 2005); possibly because
254 the species was reintroduced in a protected area without natural predators, with temperate climatic conditions (Toïgo et
255 al. 1997; Scillitani 2011) and without domestic flocks.
256
257 This increase is related to the increase in the area occupied by the species, initially only in relation to the main core
258 (core area), but later in a general way in recent years. As suggested by Vander Wal et al. (2014), saturation of
259 encounters and avoidance suggests that the benefits of social behaviour may become costs when two individuals share
260 a large extent of their home range. Besides, individuals compete over resources within a shared space (Rieucau and
261 Giraldeau 2011). On the other hand, a higher density could cause an increase in aggressive interactions as occurs with
262 other gregarious ungulates (Weckerly 1999). These factors could force the dispersion of some specimens and enhance
263 the rate of colonisation of new territories (Fuller 2007), as has happened with other ungulate populations (Messier et al.
264 1988).
265
266 Our results suggest that the population is properly settled in the area of reintroduction, as has also been found by other 267 authors (Messier et al. 1988), at least until theyDraft reach excessive densities. In this situation, there is a low dispersion of 268 the specimens that results in an excessive concentration in very specific areas, which is a factor that negatively affects
269 the vegetation (Refoyo et al. 2014) This low dispersion matches with the indicated by Pedrotti (1995) for the Alpine ibex
270 in the Alpi Orobie or for other ungulates as the roe deer ( Capreolus capreolus ) in mediterranean areas (Rossell et al.,
271 1996; Calenge et al., 2005).
272
273 The benefits of social behaviour may become costs (Vander Wal et al. 2014) when the densities reach very high values
274 (44 ind./km 2). For populations of group-living species, social structure likely has important demographic consequences
275 and can lead to dynamics that are qualitatively different from those of homogeneous populations (Bateman et al. 2012)
276 and that could force the species to faster dispersion. This pattern of increase to high density followed by expansion into
277 a new range is similar to that described for other ungulates (Larter et al. 2000).
278
279 It was found that the differential use of the different habitats present in the study area occurred depending on the time of
280 year, coinciding with what has been found for the species in the populations located in the Sierra de Gredos and Sierra
281 Nevada, where grasses are eaten more regularly in spring and early summer, but as the year progresses, woody
282 species gain prominence (Martinez 2002). Something similar happens in Puertos de Tortosa-Beceite, where woody
283 plants, in all seasons, are consumed in greater quantities than herbaceous plants, with emphasis in winter (88% of the
284 diet). Herbaceous plants are consumed more in spring and early summer (Martínez 1994). Petrotti (1995) indicates that
285 winter ranges characterize ibex as an insular species, especially by females and tend to remain in their traditional
286 ranges while increasing their density (Gauthier et al. 1994).
287
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288 Overall, in areas with abundant grasses (Gredos and Sierra Nevada) the species feeds on them, leaving woody plants
289 for winter when snow forces animals to a lower altitude. However, in areas with abundant woody plants (oak, pine, etc.)
290 the species feeds mainly on them (Cazorla, Beceite, and Tortosa), leaving grasses for times where these are abundant
291 (spring).
292
293 Differences detected between the availability and use of different habitats by the species may be due to the availability
294 of different trophic resources. As has been suggested in many studies, weather conditions are factors that affect the
295 processes of seasonal migration of many wild ungulates, mainly due to trophic resources (Gaillard et al. 2000; Clutton-
296 Brock and Coulson 2002; Garrott et al. 2003; Jacobson et al. 2004; Wang et al. 2006). Range expansion may delay
297 responses to food limitation, such as diminished survival and fecundity, until new areas can no longer be colonised to
298 provide additional forage (Messier et al. 1988; Larter et al. 2000), forcing the population to migrate (Fuller et al. 2007).
299 Ungulate populations generally become more sensitive to density-independent factors that affect resource availability as
300 they approach high densities (Saether 1997; Gaillard et al. 1998, 2000).
301
302 Finally, the high density of ungulates can cause excessive pressure on vegetation (Palacios et al. 1989; Álvarez and
303 Ramos 1991; Perea et al. 2014). High densities of ibex in a protected area may threaten certain endangered plant 304 species (Perea et al. 2015), so continuous monitoriDraftng of the population is a very effective tool to know the evolution of it 305 and establish the control measures that are necessary to reduce the population to environmentally acceptable levels.
306
307 The Iberian ibex population increase has been the cause of changes in the fauna and flora of the National Park of Sierra
308 de Guadarrama. Our data allow us to demonstrate the importance of monitoring reintroduced populations and
309 establishing the need to control the reintroduced populations to avoid these alterations in a protected area. It is
310 necessary to properly manage reintroduced populations in order to allow the coexistence of the different species in
311 protected areas.
312 Protected areas are essential for the reintroductions of Iberian ibex because they improve the probability of success
313 (Pérez et al, 2002; Moço et al., 2006; Carnevali et al., 2009; Goldstein and Rominger 2011; Refoyo et al., 2012),
314 however, as it is demonstrated in this work, the new reintroductions should consider the low dispersion of the specimens
315 as their seasonal distribution to reduce effects on threatened plant species. It is also essential to know the capacity of
316 the environment and the establishment of control measures to prevent the population to acquire densities that the
317 environment can not support.
318 Acknowledgments
319 This study was possible thanks to the collaboration of those in charge of the former Regional Park of the Cuenca Alta
320 del Manzanares, now the National Park of the Sierra de Guadarrama, and the technical staff of the company Estudios
321 Territoriales Integrados, S.L., who collaborated with us in data collection .
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493 Table 1: Available area in ha and % of total surface of the National Park for each 494 established habitat and Total area. We group the habitat into nine different vegetation 495 types (Table S1) (wet forest, thermophilic forest, rain forest, wooded areas, wet shrub, 496 thermophilic shrubs, wet grassland, thermophilic grassland, and rocky areas with 497 vegetation).
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 Draft 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
4 98
499
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500 Table 2. Evolution of the number of animals and density of Iberian ibex (Capra 501 pyrenaica , Schinz, 1838) between 2000 and 2014 in Sierra de Guadarrama National 502 Park. KIAs. Abundace index; f(0) value of probability density function at zero for line 503 transects = 1/ u (u = W*p; W = width of line transect; p = probability of observing an 504 object in defined area. Truncating distance (m) = maximum detection distance.
Year Number KIAs Density Coefficient Truncating Interval f(0) p Degrees 95% confidence of (ind/km 2) of variation distance (m) numbers of interval animals (ind/km) (CV%) freedom 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 Draft
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507 Table 3: Occupied area (ha) by 50%, 75% and 95% of the species distribution using the 508 Kernel distribution by years (in parenthesis monitoring season).
509
Ocuppied area (ha) Field seasons 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
5 10
511 Draft
512
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513
514 Table 4: Total available area by Habitat present in the National Park and available area 515 by Habitat used by ibex (Capra pyrenaica , Schinz, 1838) populations in each seasons 516 (in ha and %).
Habitat used Habitat used Habitat used Available area Habitat Spring Summer Autumn
Ha % Ha % ha % ha %
Wet forest 14.2 0.18 0 0 0 0 0 0
Thermophilic 873.1 11.24 46.77 5.16 4.00 0.21 56.87 7.45 forest
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 496.9 6.39 26.10 2.88 13.51 0.71 28.93 3.79 scrub Draft
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 55.5 0.71 0 0 0 0 0 0 grassland
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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519 Figure 1: Study area with the boundaries of the National Park of Sierra de Guadarrama; 520 Samples area, Transects and Reintroduction point.
521 Figure 2: Occupied area for each field seasons included 50%, 75% and 95% of the 522 population by Kernel distribution The occupied area is represented in gradient gray with 523 darker maximal values (95%). and clearer lower (50%). It is also indicated the 524 reintroduction point.
525 Figure 3: Relationship between abundance index (KAI,s) in no reproductive field 526 seasons and occupied area by 50, 75 and 95% of the population by Kernel distribution.
527 Figure 4: Relationship between Field seasons and distance form releasing point found in 528 each field seasons and the extent of the area occupied by the species within the National 529 Park (post-hoc Scheffé test: MS = 3003E3, df = 557; p < 0.001 (Distance/field seasons)
530
531 Draft
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532 Figure 1: Study area with the boundaries of the National Park of Sierra de Guadarrama; 533 Samples area, Transects and Reintroduction point.
Draft
534
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535 Figure 2: Occupied area for each field seasons included 50%, 75% and 95% of the 536 population by Kernel distribution The occupied area is represented in gradient gray with 537 darker maximal values (95%). and clearer lower (50%). It is also indicated the 538 reintroduction point.
Draft
539
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540
541 Figure 3: Relationship between abundance index (KAI,s) in no reproductive field 542 seasons and occupied area by 50, 75 and 95% of the population by Kernel distribution.
Field seasons
2000 2003 2007 2014 14000
Ika;(core area 50%): r 2 = 0,9783; r = 0,9891; p = 0,0109 12000 Ika; (core area75%): r 2 = 0,9303; r = 0,9645; p = 0,0355 Ika; (core area 95%): r 2 = 0,7785; r = 0,8823; p = 0,1177
10000
8000
6000 Occupied area (ha) 4000 Draft 2000
50% 0 75% 3 4 5 6 7 8 9 95% 543 KAI,s 544
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5 45
546 Figure 4: Relationship between Field seasons and distance form releasing point found in 547 each field seasons and the extent of the area occupied by the species within the National 548 Park (post-hoc Scheffé test: MS = 3003E3, df = 557; p < 0.001 (Distance/field seasons)
Current effect: F(5, 557)=10,845, p=,00000 Vertical bars denote 0,95 confidence intervals 5000
4500
4000
3500 DISTANCE (m.) DISTANCE 3000
2500 Draft
2000 2000 2003 2005 2007 2010 2014 Field seasons 549
25 https://mc06.manuscriptcentral.com/cjz-pubs 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
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 Wet Wet forest TOTAL 873.1 Mixed pine forests of an adult repopulation of Pinus pinaster and P. nigra with 25%–50% of rock outcropping; >50% pine 195.4 cover with brushwood Mixed pine forests of Pinus pinaster and an adult repopulation of P. nigra with 5%–25% of rock outcropping; 20%–50% 5.2 pine cover with brushwood
Mixed natural pine forests of Pinus pinaster, P. sylvestris and P.nigra with 5%–25% of rock outcropping; >50% pine cover 2.7 with brushwood 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
Thermophilic forest Thermophilic 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
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
Rain forest Rain 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
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 Wooded areas Wooded Rocky area with cover grade >50% with trees, 5%–20% brushwood with Pinus sylvestris and Pinus uncinata. 178.4 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 15 oaks and ferns
Rocky area with cover grade >50% without wooded thicket, with brushwood and rockroses 217.7 Thermophilic scrub Thermophilic 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 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 Wet Wet brushwood 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 TOTAL 312.5
Cervum grazes (Nardus stricta) and wet grasslands with rock outcropping; 5%–25% pure with scrub and /or ferns and 103 brooms
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 Wet Wet grassland 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
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
Xerophilous grassland Xerophilous Xerophytic grassland with rock outcropping; 5%–25% pure with lavenders and pure with juniper 4.2
TOTAL 14.6
Sports areas 5.2 Family houses and townhouses 5.8
Urban areas and urbanized areas 2.7 Urban areas Urban Other (camping, cemeteries,…) 0.9 OVERALL TOTAL 7770.2