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The successful introduction of captive bred takhi or Przewalski’s horse, Equus ferus przewalskii, into Mongolia in the 1990s is a good example of the benefits of ex situ conservation and one of the few examples of the recovery of an animal after it became extinct in the wild. This is also particularly interesting because virtually nothing was known about how takhi lived before they died out, and the introductions have enabled us to study how they have settled, and their ecology and behaviour within their former natural range. In this paper, we describe the movement, home range size and shape, and habitat use of takhi at one of the release areas, the 570 km2 Hustai National Park in Mongolia. Harem home ranges varied between 129 and 2399 ha, with 80% core areas of between 61 and 1196 ha. There was no relationship between range size and harem size, or length of time since release. Initially, harems stayed near their release enclosures, but over time they established home ranges further away. There was little overlap between home ranges of different harems, but neither was there evidence of exclusive range use. The more nutritious vegetation at lower elevations was preferentially selected. Thus the present situation looks good, but, as the population continues to grow, we anticipate that there will be potential problems related to intraspecific competition for water and vegetation resources, and the potential for hybridisation with domestic horses belonging to the local people. We consider the time it may take for takhi to reach carrying capacity within Hustai National Park and emphasise that continual monitoring of the population is essential because interventional management is likely to be required in the future.
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Habitat use and spatial dynamics of takhi introduced to
Hustai National Park, Mongolia
Sarah R.B. King
1
, John Gurnell
*
School of Biological Sciences, Queen Mary, University of London, Mile End Road, London E1 4NS, UK
Received 27 August 2004
Abstract
The successful introduction of captive bred takhi or PrzewalskiÕs horse, Equus ferus przewalskii, into Mongolia in the 1990s is a
good example of the benefits of ex situ conservation and one of the few examples of the recovery of an animal after it became extinct
in the wild. This is also particularly interesting because virtually nothing was known about how takhi lived before they died out, and
the introductions have enabled us to study how they have settled, and their ecology and behaviour within their former natural range.
In this paper, we describe the movement, home range size and shape, and habitat use of takhi at one of the release areas, the 570 km
2
Hustai National Park in Mongolia. Harem home ranges varied between 129 and 2399 ha, with 80% core areas of between 61 and
1196 ha. There was no relationship between range size and harem size, or length of time since release. Initially, harems stayed near
their release enclosures, but over time they established home ranges further away. There was little overlap between home ranges of
different harems, but neither was there evidence of exclusive range use. The more nutritious vegetation at lower elevations was pref-
erentially selected. Thus the present situation looks good, but, as the population continues to grow, we anticipate that there will be
potential problems related to intraspecific competition for water and vegetation resources, and the potential for hybridisation with
domestic horses belonging to the local people. We consider the time it may take for takhi to reach carrying capacity within Hustai
National Park and emphasise that continual monitoring of the population is essential because interventional management is likely to
be required in the future.
Ó2005 Elsevier Ltd. All rights reserved.
Keywords: Home range; Habitat use; Equus przewalskii; Takhi; Reintroduction
1. Introduction
Collections of animals and plants are increasingly
being seen as opportunities for ex situ conservation,
notably with respect to captive breeding programmes
and subsequent release of captive-bred animals back in
to the wild. However, to date there have been few suc-
cessful examples of such a strategy; approximately
27% of the 116 reintroductions considered by Fischer
and Lindenmayer (2000) were classified as failures, with
the success of a further 47% not known. Moreover, the
respective roles of ex situ and in situ conservation strat-
egies are open to debate (e.g. Balmford, 2000; Entwistle
and Dunstone, 2000). Captive breeding and reintroduc-
tions are lengthy, complex and expensive processes
(IUCN, 1995; Balmford, 2000) but may be vital for
the survival of a species in the wild (e.g. Spalton et al.,
1999; Kleiman and Rylands, 2002).
Before reintroduction of captive bred takhi or Prze-
walski horses, Equus ferus przewalskii, into Mongolia
in the 1990s, the last authenticated sighting of a takhi
in the wild was in 1969, near the Tachyn-shar mountains
0006-3207/$ - see front matter. Ó2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2005.01.034
*
Corresponding author. Tel.: +44 20 7882 3041.
E-mail addresses: kingsrb@yahoo.com (S.R.B. King), j.gurnell@
qmul.ac.uk (J. Gurnell).
1
Present address: School of Renewable Natural Resources, Univer-
sity of Arizona, Biosciences East, Rm. 325 Tucson, AZ 85721, USA
www.elsevier.com/locate/biocon
Biological Conservation 124 (2005) 277–290
BIOLOGICAL
CONSERVATION
in the western Gobi desert (Bouman and Bouman,
1994). The captive population of takhi in European
and North American animal collections is descended
from animals captured at the beginning of the 20th Cen-
tury. In 1945, only 31 horses remained in captivity, and
only nine of these bred. A species survival plan was set
up in the USA in 1979, followed in 1986 by a European
breeding programme (Europaisches Erhaltungzucht-
Programm or EEP; Bouman and Bouman, 1994). By
the start of the 1990s, there were more than 1500 horses
in captivity. In 1992, reintroductions began to Tachyn
Tal, by the German Christian Oswald Foundation and
the Mongolian government, and to Hustai National
Park, by the Dutch Foundation Reserves Przewalski
Horse in association with the Mongolian Association
for the Conservation of Nature and the Environment
(Bouman, 1998).
For animals kept in captivity for many generations,
there is a potential for loss of genetic variability (Foose,
1986; Ryder, 1986; May, 1991) or attenuation in sur-
vival skills (Box, 1991; Brock et al., 1994; Shepherdson,
1994; McPhee, 2003; Mathews et al., 2005). Thus there
was concern that released takhi would not be able to
survive after being bred in captivity for 13 generations
(Klimov and Orlov, 1982). Moreover, there was no
knowledge about the ecology of takhi in the wild before
they became extinct, and so it was not clear how the re-
leased animals would cope in their new surroundings.
Between 1995 and 2000, we carried out a series of
studies on the behaviour and ecology of released takhi
in Hustai National Park. In this paper, we report on
the spatial dynamics and habitat use of the horses.
Against the background of changes in population size
during the study period, we consider the establishment
of home ranges by the harems in relation to their release
point, size, interrelationship between the ranges of dif-
ferent harems, and utilisation of the habitat within these
ranges. Finally, we consider what might happen in the
future as the population continues to increase. By exam-
ining these factors we hope to be able to provide infor-
mation on a reintroduction that currently appears
successful, with an aim to helping future reintroduction
projects.
1.1. Study site
Hustai National Park is 150 km south west of Ulaan
Baatar, the capital city of Mongolia. It covers 570 km
2
along a southwestern spur of the Khentei range of
mountains (47°410N, 105°540E). It was made a reserve
in 1993 and designated a National Park in 1999. Hustai
National Park borders the Tuul River, with an altitude
of between 1100 and 1842 m and a continental climate
(mean annual temperature = 0.2 °C and yearly rain-
fall = 270 mm, most of which falls in the summer; Wallis
de Vries et al., 1996). During the study, horses were ob-
served in temperatures of between 14 and +34 °C.
Hustai National Park has a mountain forest steppe hab-
itat that consists of steppe, meadow, grassland, shrub
and woodland communities (Wallis de Vries et al.,
1996). Water flows down most of the major valleys as
streams, which are permafrost fed and covered by gravel
at places along their length.
1.2. Study animals
Between 1992 and 2000, 84 horses were brought to
Hustai National Park from reserves in Europe, where
they had been kept in large (over 40 ha) grassy enclo-
sures. The horses were released as a harem into acclima-
tisation enclosures of 40–45 ha; these enclosures were a
minimum of 4 km apart and visually separated from
each other. Although most of the horses had been kept
together in Europe and were released together, others
were introduced on arrival in Mongolia. Harems are re-
ferred to by the name of the dominant stallion (see Table
1). One harem was given a hard release (TurgenÕs har-
em), but other releases followed a 6–24 month acclima-
tisation period. Up to four harems other than the ones
studied were also present towards the end of the study
period (see Table 1).
2. Methods
Mongolian rangers and biologists gathered positional
data of the harems at Hustai National Park for the full
year from 1995 to 1997, and for the winter months
(December to March) from 1998 to 2000. Detailed
observations were carried out by the first author be-
tween June 1998 and July 2000, totalling 860 h. The
standard procedure was to find a harem at dawn and
follow it until 1400 h. The same harem was located at
1400 h the next day and followed until dusk. Every
10 min during an observation session the position of
the harem was recorded with a Global Positioning Sys-
tem and marked on a large-scale map, together with
weather details. Once accustomed to the presence of
the observer, the horses were watched on foot from a
distance of about 15 m and their behaviour recorded;
these results will be considered elsewhere (see King,
2002).
The per capita growth rate per year for harems with
more than three years of census data have been esti-
mated using the formula r= [ln(N
t
/N
0
)]/t, where N
t
is
the number of animals alive tyears after reintroduction,
and N
0
is the number of animals released (Komers and
Curman, 2000). Where mares changed harems, for the
purposes of this calculation they were included in the
harem with which they were released. However, because
mares move between harems, and the number of horses
per harem is likely to decrease over time as the number
278 S.R.B. King, J. Gurnell / Biological Conservation 124 (2005) 277–290
Table 1
Harem composition and harem range and core areas (ha) in Hustai National Park between 1994 and 1998
Harem Year Release
enclosure
No. adults
and juveniles
No. foals Total
harem
size
No. fixes Period Home range Core area
Paritet 1995 1 7 1 8 142 July–December 191 96
1996 7 2 9 194 January–December 751 275
1997 6 3 9 131 January–August 881 257
1998
*
7 4 11 423 June–October 1233 499
1999
*
11 5 16 1166 April–November 1223 408
2000
*
14 2 16 586 May–June 709 126
Mean (SD) 832 (387) 277 (156)
Bayan 1998
*
1 6 2 8 395 June–October 210 61
1999
*
8 2 10 628 April–October 1114 496
2000
*
9 3 12 315 May–June 748 320
Mean (SD) 690 (455) 292 (219)
Margad 1999
*
a 14 2 16 762 May–October 684 345
2000
*
12 3 15 270 May–June 277 126
Mean (SD) 480 (288) 235 (155)
Khaan 1994 1 6 4 10
1995 9 3 12 194 April–December 1089 469
1996 9 4 13 228 January–December 999 367
1997 13 5 18
1998
*
15 7 22 389 June–October 609 249
1999
*
7 4 11 199 April–June 1100 523
Mean (SD) 949 (231) 402 (121)
Ares 1998
*
2 6 3 9 144 July–October 744 240
1999 9 0 9
2000 9 3 12
Patron 1994 2 6 5 11
1995 9 3 12 159 April–December 2399 1196
1996 14 4 18 156 January–December 1904 894
1997 13 2 15 141 January–July 1653 718
1998 12 7 19
1999 15 5 20
2000 16 5 21
Mean (SD) 1985 (380) 936 (242)
Turgen 1996 b 4 1 5 96 June–December 751 260
1997 3 0 3 6 January/April 129 68
Mean (SD) 440 (440) 164 (135)
Bohemian 1998 4 6 0 6
1999 3 0 3
Mark 1998 5 7 0 7
Manlai 1999 a 4 0 4
Mangir 1999 a 4 2 6
2000 6 1 7
Harem range area = 95% kernel estimates, core area = 80% kernel estimates. a = wild born bachelor stallion acquired mares. b = harem hard released
from travelling crates near release enclosure 1. There was no difference in mean home range size or mean core area among years, irrespective of harem
(home range: F
5, 15
= 0.56, P= 726; core area: F
5, 15
= 0.78, P= 0.578; these include the data for Ares). Mongolian rangers collected data prior to
1998.
*
= data collected by SRBK; = no data.
S.R.B. King, J. Gurnell / Biological Conservation 124 (2005) 277–290 279
of stallions competing for mares increases (Linklater
and Cameron, 2000; Linklater et al., 2000), changes in
numbers of individuals within harems do not reflect
overall population growth. Thus, we also consider pop-
ulation growth based on changes in population numbers
between 2002 and 2004 (see Anonymous, 2004); no fur-
ther introductions took place after 2001.
Home range and habitat use data were analysed using
Arcview 3.2. One hundred per cent and 95% kernel har-
em range areas were calculated (see Worton, 1989; Sea-
man and Powell, 1996; Hooge and Eichenlaub, 1997),
the latter excluded occasional forays outside the areas.
Utilisation curves were used to estimate core areas; in
all cases these were 80% of the total range area (King,
2002).
ANOVA was used to test whether the yearly range
areas since release changed in a systematic way. Param-
atric or non-paramatric correlation techniques were
used as appropriate to compare the area overlap of each
yearly harem range with the range in its previous year,
and the movement of range centres from one year to
the next. Harem range use was examined according to
season and temperature using Kruskal–Wallis tests, be-
cause the data were not normal. For analyses of sea-
sonal and altitudinal effects on activity, movement,
and range size, the yearly range or core areas for partic-
ular harems have been treated as independent.
Eleven vegetation categories or habitat types were
distinguished within Hustai National Park (Wallis de
Vries et al., 1996). The areas covered by the different
habitat types within each home range, and those that
were grazed or used for shelter or water, were measured,
and habitat selection indices (Manly et al., 1993) calcu-
lated. Preferences were formulated using v
2
tests. The
seasonal (spring – March to May, summer – June to Au-
gust, autumn – September to November, and winter –
December to February) use of vegetation was based
on data from all years and this was compared with the
vegetation available to them within their total range
area over all years.
3. Results
3.1. Numbers and density of horses
Of 115 foals born between 1994 and 2001 (Table 1),
57% were still alive at the end of the study reported here
(see Anonymous, 2004, for an update of population
numbers). Twenty-six per cent of deaths were caused
by babesiosis (piroplasmosis), which particularly af-
fected newly released horses (three died from this in
1998 and nine in 1999; previous deaths from this disease
are also possible, but were not diagnosed). Most other
deaths were foals (63% of all deaths). In each year
(1995–2000), on average, foals made up 25% of the pop-
ulation, and 12% of the population died. The increase in
births in 1998 was mostly produced by three free-rang-
ing harems (19 of the 25 births). Predation by wolves,
Canis lupus, was low with a mean of 2.8 horses taken
each year, all one year old or less (16% of all foals born
between 1994 and 1999). It appears unlikely that preda-
tion affected population growth at the time of the study.
In July 2000 there were 91 free ranging horses, a
density of 0.2 horses km
2
over the entire Park. The
yearly mean density of horses within a harem between
1995 and 2000 was 1.8 horses km
2
(N= 24 harems,
SD = 1.3 horses km
2
, range from 0.5 horses km
2
for Patron in 1995 to 5.4 horses km
2
for Margad
in 2000). In 1995, 1996 and 1998 the home range
and size of every harem was known. The density of
horses over the area covered by these harems was
0.8 horses km
2
in 1995, 1.2 horses km
2
in 1996
and 1.8 horses km
2
in 1998. In 1999, density of har-
ems within Hustai National Park was 0.01 har-
ems km
2
(N= 7).
The per capita growth rates (r) of three reintroduced
harems with census data spanning more than three years
were: Paritet 0.69 – 1995 to 2000, Patron 0.65 – 1994 to
2000, and Khaan 0.79 – 1994 to 1998. Thus, these har-
ems were very successful and doubled in size over these
time periods. However, Khaan lost half his harem in
1999 to Margad when numbers dropped from 22 to 11
(Table 1), and he subsequently lost the rest of his harem
the year after. A more realistic indication of population
growth rate can be obtained by considering population
numbers after 2001, when no further introductions took
place. In 2002 there were 142 horses in Hustai National
Park, and this increased to 162 horses by June 2004, of
which 84 were reintroduced (Anonymous, 2004). This
gives an overall per capita growth rate (r) of 0.066 per
year. This growth rate must be treated with caution
since it is based on only three years of data.
3.2. Harem range and core areas
Between 1995 and 2000, 95% kernel harem range
areas ranged from 1.3 to 24.0 km
2
and 80% core use
areas from 61 to 499 ha (Table 1,Fig. 1). There was
no difference in range area among years (F
5,15
= 0.56,
P= 0.726; note, horses were observed for different peri-
ods of time within each year). Both mean range and core
areas were significantly different among harems (range
area: F
5,14
= 6.61, P= 0.002; core area: F
5,14
= 7.65,
P= 0.001, these exclude the single home range estimate
for Ares, Table 1); mainly because PatronÕs mean range
and core areas were significantly larger than those of the
other harems (Table 1).
There was no correlation between number of fixes
and size of harem range or core areas (Table 1; range
area: r
s
= 0.035, N= 21, P= 0.881; core area:
r
s
= 0.092, N= 21, P= 0.691). In addition, there was
280 S.R.B. King, J. Gurnell / Biological Conservation 124 (2005) 277–290
no correlation between number of months since release
and harem range (r= 0.22, N= 31, P= 0.235) and core
area (r=0.01, N= 31, P= 0.976), or between the
number of adults and juveniles in each harem and range
(r
s
= 0.220, N= 21, P= 0.337) or core area (r
s
= 0.346,
n= 21, P= 0.124). Despite the small sample sizes, we
tested for a year effect, but there was no significant dif-
ference in range or core area among years (range area:
H
5
= 3.56, P= 0.615; core area: H
5
= 5.09, P= 0.404).
3.3. Harem range shifts between years
Overlap of individual harem ranges between years
varied between 31% and 100% (mean = 59%, N= 11,
SD = 26%) (Table 2). Overlap of yearly core areas var-
ied between 11% and 100% (mean = 45%, N= 11,
SD = 27%; Table 2). There was no significant correla-
tion between the number of years since release and the
degree of overlap (range area r
s
=0.34, N= 11,
P= 0.313; core area r
s
=0.40, N= 11, P= 0.255;). In
general the trend was for a decrease in range overlap
over the years (Table 2).
The harem range centres did not move very far from
one year to the next, with a minimum movement of
0.4 km and a maximum of 1.8 km (Table 3). However,
distance from initial range centre after release to range
centres the following years appear to show a movement
away from the release enclosure. The harems tended to
stay close to their acclimatisation enclosures in the year
after release, then move further away to settle in a near-
by valley.
3.4. Overlap between the ranges of different harems
Paritet, Bayan and Khaan were released from enclo-
sure 1, Turgen from outside enclosure 1, and Ares and
Patron were released from enclosure 2. MargadÕs was
a naturally formed harem (Table 1). Overlap between
yearly harem ranges was analysed to see whether harems
moved to the same areas of Hustai National Park after
release. The degree of range overlap between harems
varied from 0% to 89% (mean = 39%, N= 21,
SD = 30%); core areas overlapped 0% to 66%
(mean = 18%, N= 21, SD = 21%) (Table 4). There was
no correlation between number of years since release
and amount of overlap between harem ranges
(r=0.26, N= 21, P= 0.253) or core areas
(r=0.17, N= 21, P= 0.459).
In 1995, when ParitetÕs harem was first released and
KhaanÕs and PatronÕs had been free ranging for one
year, there was no overlap between any of the core
areas. Core areas overlapped in 1996 after TurgenÕs
harem was released. TurgenÕs core area in 1996 and
1997 was overlapped more than any other harem except
AresÕin 1998 (Table 4b). PatronÕs core area did not
overlap with any other harem between 1995 and 1997,
and did not do so during the rest of this study (SRBK,
pers. obs.). The bachelor group moved among the home
ranges of all (SRBK, pers. obs.).
Changes in overlap among core areas indicate range
shifts and/or a change in area used. For example, Ba-
yan overlapped with Ares in 1998, but in 1999 he used
a larger area and his harem overlapped with all the
other harems. The bachelor stallion Margad took most
of KhaanÕs mares in 1999, and although they were
using the same valley, their core areas only overlapped
by 23%. By mid-summer, KhaanÕs remaining mares
were taken over by another stallion and they moved
to a different valley. Thereafter MargadÕs harem
increasingly used KhaanÕs previous range; in 2000,
50% of MargadÕs core area overlapped with KhaanÕs
1998 core area and 57% with KhaanÕs 1999 core area.
Fig. 1. Eighty percent core areas of the study harems in (a) 1998 and
(b) 1999. Ranges show the core area of the entire harem, but the names
of the dominant stallions are used. Paritet, Khaan and BayanÕs harems
were all released from enclosure 1 (in 1994, 1995 and 1998,
respectively). AresÕharem was released from enclosure 2, and Margad
acquired KhaanÕs mares in 1999.
S.R.B. King, J. Gurnell / Biological Conservation 124 (2005) 277–290 281
In 1999 and 2000 there were four other harems in
Hustai National Park (Table 1). Although there were
no data for these harems, it is possible that total overlap
was larger than shown in Table 4. The harems appeared
to distance themselves from each other but there was no
evidence of exclusive use of an area by any harem. The
harems appeared to select a home range in a valley
where there were no previous horses, and where the har-
ems were separated by ridges. However, Bayan broke
this trend by tending to use the same valley as Margad
in 2000. Moreover, observations by the Mongolian
rangers indicate that all harems sometimes use the same
valley during the winter.
3.5. Habitat use within the home range
Habitat use within the home range was influenced by
several factors including temperature, season, time of
day and the presence of flies. Although ranges were
slightly larger in winter, there was no significant differ-
ence in the size of the ranges or core areas in different
seasons (range areas: F
3,39
= 0.49, P= 0.693; core areas:
F
3,39
= 0.62, P= 0.605).
In core areas, horses were found more often at a
higher altitude during the summer than in spring or au-
tumn (there were too few data for winter; median in
summer = 1500 m, spring = 1425 m and autumn =
1400 m; Kruskall–Wallis H
2
= 613.31, P< 0.0001).
There was also a diurnal trend in the horses movements
(Fig. 2). In the morning, when it was cool, the horses
spent most of their time grazing in the valleys. As the
temperature increased, the horses moved to higher ele-
vations where they could avoid flies and direct solar
radiation by standing near rocky outcrops or in forest
until the weather cooled enough for them to come down
to graze again. This pattern was less marked in spring
Table 2
Harem home range and core area shifts expressed as a percentage of overlap from one year (column) to the previous years (row)
Year 1996 1997 1998 1999 2000
(a) Overlap of home range
Paritet
1995 100 93 75 89 87
1996 75 47 60 41
1997 62 64 45
1998 62 44
1999 37
Bayan
1998 98 18
1999 33
Margad
1999 31
Khaan
1995 78 *23 36
1996 *31 34
1997 **
1998 41
(b) Overlap of core area
Paritet
1995 100 82 24 86 89
1996 79 39 78 45
1997 48 76 44
1998 47 9
1999 25
Bayan
1998 51 0
1999 11
Margad
1999 20
Khaan
1995 61 *23 4
1996 *34 7
1997 **
1998 19
*
= no data for Khaan harem in 1997.
282 S.R.B. King, J. Gurnell / Biological Conservation 124 (2005) 277–290
and autumn when the weather was cooler and the horses
spent more time at lower elevations.
3.6. Vegetation use within the home range
Habitat composition of the different haremsÕranges
varied slightly, but all 11 vegetation classes described
by Wallis de Vries et al. (1996) were grazed at some time
(Table 5). However, all harems in all years did not ran-
domly select vegetation classes (all v
2
statistics, P< 0.05;
Fig. 3). They most frequently selected vegetation classes
from lower elevations (i.e., Meadow I, Tussock grass-
land and Lowland steppe; Fig. 3). Meadow I (domi-
nated by bentgrass, sedge and Iris lactea) was
preferred by Bayan in 1999 and 2000, and by Khaan
in 1998. Although most preferred, Meadow I was not
used by Khaan in 1999, and little by Margad.
ParitetÕs range did not include Meadow I vegetation
and they preferred Tussock grassland (Fig. 3). Tussock
grassland contains the needlegrass species Achnatherum
splendens and Stipa krylovii which have high levels of
crude protein and crude fibre (van Dierendonck and
Wallis de Vries, 1993). Tussock grassland was not found
in Khaan/MargadÕs home range.
Lowland steppe was selected significantly more than
most other vegetation classes, and had a high selection
index for all harems except Bayan in 1999 and Khaan
in 1998 (Fig. 3). This vegetation class includes the
wheatgrass Agropyron cristatum, which featured largely
in the diets of feral horses, and is dominated by the nee-
dlegrass Stipa krylovii.
Both Mountain steppe vegetation classes were found
in every home range, and one or both were significantly
selected by the harems (Fig. 3). These were dominated
by Festuca lenensis in Mountain steppe I and Festuca
sibirica in Mountain steppe II. Although these species
did not have the high crude protein content of needle-
grass they had a higher crude fibre content (van Diere-
ndonck and Wallis de Vries, 1993).
Grazing was observed in Woodland by most harems
in most years (Fig. 3) and understorey vegetation such
as Spiraea media and Cotoneaster melanocarpa was ea-
ten rather than trees. The area covered by other vegeta-
tion classes (Upland steppe, Shrubland I, Shrubland II,
Meadow II and Scrub) was small.
3.7. Seasonal use of the vegetation
The harems selected more vegetation classes in spring
and autumn than in summer, and few classes were se-
lected in winter (Fig. 4). From spring to autumn vegeta-
tion classes at mid to low elevations were used, with the
exception of using woodland in the spring. In winter,
vegetation at mid elevations was used. Lowland steppe
was selected in every season of the year, and Shrubland
I and Mountain steppes I and II were selected in every
season except winter. Harems observed in winter pre-
ferred Lowland steppe. Woodland was not used at all
in the autumn and winter, and Tussock grassland and
Upland steppe were selected by at least two of the three
harems in spring and autumn, but not in summer.
4. Discussion
4.1. Numbers
The reintroduced population of takhi increased in
size during the study period, and continues to grow
(Anonymous, 2004). This was despite loss of half the
Table 3
Distance from the centre of the home range in one year to the centre in all other years since release
Year 1996 1997 1998 1999 2000
Paritet
1995 0.5 1 2.7 1.6 1.6
1996 0.5 2.3 1.2 1.2
1997 1.8 0.8 0.8
1998 1.2 1.2
1999 0.1
Bayan
1998 0.5 3.2
1999 2.7
Margad
1999 1.4
Khaan
1995 0.4 *1.6 2 2.8
1996 *1.3 1.7 2.5
1997 ***
1998 0.9 >1.2
The arithmetic mean of all points was used to find the centre. Figures given are the distance from the centre of the row to the centre of the column.
*
= no data for Khaan harem in 1997.
S.R.B. King, J. Gurnell / Biological Conservation 124 (2005) 277–290 283
Table 4
Overlap of home ranges and core areas between harems expressed as a percentage of the area covered in each year
Year Harem Home range area (ha) Harem Total % of core area overlapped
Paritet Khaan Patron
(a) Home range
1995 Paritet 191 86 0 86
Khaan 1089 15 4 19
Patron 2399 0 39 39
1996 Khaan Paritet Turgen Patron
Khaan 999 42 25 0 51
Paritet 751 56 53 0 87
Turgen 751 34 53 0 85
Patron 1904 0 0 0 – 0
1997 Paritet Turgen Patron
Paritet 881 13 0 13
Turgen 129 89 0 89
Patron 1653 0 0 0
1998 Khaan Paritet Bayan Ares
Khaan 609 – 14 0 29 35
Paritet 1233 7 0 12 14
Bayan 210 0 0 41 41
Ares 744 24 19 12 18
1999 Paritet Bayan Margad Khaan
Paritet 1223 8 6 37 40
Bayan 1114 9 – 11 9 23
Margad 684 10 19 49 66
Khaan 1100 41 9 30 – 68
2000 Paritet Bayan Margad
Paritet 709 8 0 8
Bayan 748 8 5 13
Margad 277 0 14 14
(b) Core area
Paritet Khaan Patron
1995 Paritet 96 0 0 0
Khaan 469 0 0 0
Patron 1196 0 0 0
1996 Khaan Paritet Turgen Patron
Khaan 367 5 20 0 21
Paritet 275 7 – 34 0 35
Turgen 260 29 36 0 66
Patron 894 0 0 0 – 0
1997 Paritet Turgen Patron
Paritet 257 16 0 16
Turgen 68 59 0 59
Patron 718 0 0 0
1998 Khaan Paritet Bayan Ares
Khaan 249 – 4 0 0 5
Paritet 499 2 0 3 5
Bayan 61 0 0 33 33
Ares 240 0 5 8 – 14
1999 Paritet Bayan Margad Khaan
Paritet 408 5 0 31 35
Bayan 496 4 5 1 9
Margad 345 0 7 23 32
Khaan 523 0 1 15 – 39
2000 Paritet Bayan Margad
Paritet 126 0 0 0
Bayan 320 0 1 1
Margad 126 0 1 1
Columns overlap rows. There was no difference among the harems in the overlap of home ranges (F
6,14
= 1.66, P= 0.204), but there was a difference
among the overlap of core areas (F
6,14
= 3.53, P= 0.024).
284 S.R.B. King, J. Gurnell / Biological Conservation 124 (2005) 277–290
horses released in 1998 to babesiosis. Babesia equi and
B. caballi are endemic to Mongolia and transmitted by
the tick Dermacentor nutalli. Unless treated, between
25% and 100% of cases are fatal (West, 1992). However,
animals can acquire immunity during the course of an
infection, and mortality depends on the general immune
status of the animal, and the virulence of the piroplasm
(Schein, 1988). Foals can be infected during pregnancy,
but gain protection from maternal antibodies (Donnelly
et al., 1982). Thus local domestic horses and takhi born
in Mongolia do not appear affected by the disease. In
contrast, newly introduced animals with no immunity
and that may become stressed for reasons such as trans-
port, release, or poor body condition in early spring, are
likely to be susceptible. In particular there is a high mor-
tality rate during the initial infection of older animals
Hour of day
4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21
Elevation (m)
1250
1300
1350
1400
1450
1500
1550
Temperature (C)
0
5
10
15
20
25
Grazing
Resting
Moving
Mean elevation
Mean temperature
Fig. 2. Diurnal activity of the horses (% of total) according to mean elevation and temperature (SDs not included). The horses were observed in
different activities at different times of day (H= 17.38, d.f. = 4, P< 0.002), and at different temperatures (H= 577.12, d.f. = 4, P< 0.0001) and
different elevations (H= 630.21, d.f. = 4, P< 0.0001). The horses were found at higher elevations in the middle of the day (H= 120.35, d.f. = 15,
P< 0.0001) and at higher temperatures (H= 1279.18, d.f. = 15, P< 0.0001).
Table 5
Habitat classes at Hustai National Park according to landscape features and dominant plant species (based on Wallis de Vries et al., 1996)
Habitat class Dominant species Landscape Altitude Slope
Scientific name Common name
Meadow I Iris lactea Dwarf iris Streamside 1100–1400
Tussock grassland Achnatherum splendens Needlegrass Valley terrace 1100–1400
Lowland steppe Artemisia adamsii Wormwood Footslope 1100–1400 South
Stipa krylovii Needlegrass
Upland steppe Thermopsis lanceolatus False lupin Footslope 1300–1500 South
Stipa krylovii Needlegrass
Shrubland I Caryopteris mongholica Bluebeard Rocky slope 1300–1600 South
Amygdalus pedunculata Sweet almond
Mountain steppe I Festuca lenensis Tundra fescue Ridge & topslope >1300
Mountain steppe Festuca siberica Siberian fescue Mountain slope >1400 North
Shrubland II Spiraea aquilegifolia Spiraea Gully 1100–1600
Meadow II Geranium pratense Meadow cranesbill Combe >1300
Woodland Betula platyphylla Asian white birch Mountain slope >1400 North
Scrub Betula fusca Birch Topslope >1400 North
S.R.B. King, J. Gurnell / Biological Conservation 124 (2005) 277–290 285
when brought from a country, such as Holland, where
B. equi is not endemic (Schein, 1988).
Most losses to the population resulted from the death
of foals. Diseases such as strangles (infection by Strepto-
coccus equi) accounted for some of these deaths, as well
as predation and the harshness of the Mongolian winter.
4.2. Home range use
Harem range sizes varied between 1 and 24 km
2
(mean 9.2 km
2
,N= 21, SD = 5.64 km
2
) and are similar
in range to feral horse populations in Alberta, Canada
(2.6–14 km
2
;Salter and Hudson, 1982) and in New Zea-
land (1–18 km
2
;Linklater et al., 2000), but smaller than
those in North America (8–48 km
2
;Pellegrini, 1971; Fe-
ist and McCullough, 1976; Berger, 1977, 1986; Miller,
1983a). Home ranges of horses in England (2.5–
3.2 km
2
;Tyler, 1972; Gates, 1979), and barrier islands
(3–6 km
2
;Keiper, 1976; Zervanos and Keiper, 1979;
Rubenstein, 1981) were generally smaller than at Hustai
National Park. In general, differences in home range size
are related to vegetation quality and distribution of re-
sources such as water and shelter (Leuthold, 1977). No
trends in harem range size with time after release were
found, or with harem size (cf. Zervanos and Keiper,
1979; Berger, 1986; Linklater et al., 2000), and there
was no suggestion that forage was limiting. However,
Mongolia suffers from periodic droughts and severe win-
ters and these will affect the vegetation growth. It is pos-
sible that harem ranges will be smaller when forage
quality is good; this was found in reintroduced Arabian
oryx, Oryx leucoryx, whose ranges became smaller in
areas where rain had fallen (Corp et al., 1998).
Similarity in size of reintroduced takhi home ranges
and feral horses under similar circumstances implies
not only that feral horse management techniques could
potentially be applied to this population, but that they
are using similar survival strategies. Thus takhi did not
Paritet harem (1998, 1999, 2000)
W
S2
MS2
MS1
S1 LS
TG
A
W
S2
MS2
MS1
S1
LS
TG
W
S2
MS2
MS1
S1
LS
TG
-1.5
-1
-0.5
0
0.5
1
0 250 500
Area of vegetation i
log wi
(a)
Khaan harem (1998, 1999)
M1
LS
US S1
MS1
MS2
M2
W
STG
LS
US
S1
MS1
MS2
S2
W
-1.5
-1
-0.5
0
0.5
1
1.5
0 150 300
Area of vegetation i
log wi
Margad (prefix 1; 1999, 2000) and Ares (prefix 2, 1998 ) harems
1M1
1LS
1US 1S1
1MS1
1MS2
1M2 1W
1M1
1LS
1US 1S1
1MS2
1M2
1W
2M1
2LS
2US
2S1
2MS1
2MS2
2M2
2W2M1
-1.4
-1.2
-1
-0.8
-0.6
-0.4
-0.2
0
0.2
0.4
0.6
0 150 300
Area of vegetation i
log wi
(c) (d)
Bayan harem (1998, 1999, 2000)
TG
LS
US
S1
MS1
MS2
S2
W
S
R
M1
TG
LS
US
S1
MS1
MS2
S2
M2
W
S
R
M1
TG
LS
US
S1
MS1 MS2
M2 W
S
-1.5
-1
-0.5
0
0.5
1
0 150 300
Area of vegetation i
log wi
(b)
Fig. 3. Habitat selection indices (w
i
, logged for presentational purposes) plotted against area of each vegetation type (i). M1 = Meadow I,
M2 = Meadow II, TG = Tussock Grassland, LS = Lower Steppe, US = Upper Steppe, S1 = Shrubland 1, S2 = Shrubland 2, MS1 = Mountain
Steppe I, MS2 = Mountain Steppe II, W = Woodland, S = Scrub, A = Agricultural Area, R = Rock and Stones. Habitat classification based on
Wallis de Vries et al. (1996). A value of log w
i
> 0 represents positive selection, = 0 represents no selection, <0 represents avoidance. Squares indicate
habitats were not selected at all, but were arbitrarily assigned a value of 0.05 so that they appear on the figure, circles indicate w
i
values significantly
different to zero (v
2
P< 0.05), triangles indicate insignificant w
i
values or where expected values <5.
286 S.R.B. King, J. Gurnell / Biological Conservation 124 (2005) 277–290
lose an ability to adapt to the wild despite 13 generations
in captivity, just as this trait has not been bred out of
horses despite 5000 years of domestication.
Characteristics of the ranges of all harems were that
they included a permanent water source, an area of
rocky outcrops near a ridge, and patches of forest that
were used in the summer. The harems of Ares, Bayan,
Khaan and Margad all overlapped at one particular
place along the stream where pools were formed and
the banks had been eroded to form a salt lick. How-
ever, in the main, the home ranges occupied separate
valleys, defined by ridges which peaked at about
1600 m, but which also joined at passes and along
watercourses.
As the size and number of harems change through
time at Hustai National Park, competition for resources
may increase. For example, larger harems have been
shown to have priority of access to water holes (Miller
and Deniston, 1979), and also tend to be found in areas
of better vegetation (Miller and Deniston, 1979; Berger,
1977, 1986). Most studies on feral horses show extensive
spatial overlap between ranges (Keiper, 1976; Miller,
1983a; Linklater et al., 2000), although horses appear
to avoid each other temporally, as they did at Hustai
National Park (King, 2002). After release, harems
stayed near the release enclosure in the first year, then
moved a little further away to settle in a valley. The har-
ems selected valleys that were visually separated from
each other, but as the population increases and more
overlap occurs, more valleys will be colonised and smal-
ler ridges within a valley may separate the harems. In
winter harems seemed more tolerant of close proximity
to each other (Mongolian rangers, pers. comm.), imply-
ing that at this time being near other harems is more
important than having exclusive access to a food source.
More work needs to be done in this season to examine if
lack of spatial segregation between harems is for ther-
mal reasons, or for greater protection against predation.
Spring
3W
3US
3TG
3S2
3S1
3S
3MS2
3MS1
3M2
3M1
3LS
2W
2US
2TG
2S2
2S1
2MS2
2MS1
2M2
2M1
2LS
1W
1US
1TG
1S2
1S1
1MS2
1MS1
1M1
1LS
-1.5
-1
-0.5
0
0.5
1
0 400 800
Area of vegetation i (ha)
log wi
Autumn
1LS
1M1
1MS1
1MS2
1S1
1S2
1TG
1US
1W
2LS
2M1
2M2
2MS1
2MS2
2R
2S
2S1
2S2
2TG
2US
2W
3LS
3M1
3M2
3MS1
3MS2
3S
3S1
3S2
3TG
3US
3W
-1.5
-1
-0.5
0
0.5
1
0
Area of vegetation i (ha)
log wi
800
Winter
1LS
1M1
1MS1
1MS2
1S1
1S2
1TG
1US
1W
3LS
3M1
3M2
3MS1
3MS2
3S
3S1
3S2
3TG
3US
3W
-1.5
-1
-0.5
0
0.5
1
0 400 800
Area of ve
g
etation i (ha)
log wi
400
(a)
(d)
(c)
Summer
3W
3US
3TG
3S2
3S1
3S
3MS2
3MS1
3M2
3M1
3LS
2W
2US
2TG
2S2
2S1
2S
2R
2MS2
2MS1
2M2
2M1
2LS
1W
1US
1TG
1S2
1S1
1MS2 1MS1
1M1
1LS
-1.5
-1
-0.5
0
0.5
1
1.5
0 800
Area of vegetation i (ha)
log wi
(b)
Fig. 4. Habitat selection indices (w
i,
, logged for presentational purposes) plotted against area of each vegetation type (i). (a) = spring, (b) = summer,
(c) = autumn and (d) = winter. First number 1 = Paritet, 2 = Bayan, 3 = Khaan/Margad. M1 = Meadow I, M2 = Meadow II, TG = Tussock
Grassland, LS = Lower Steppe, US = Upper Steppe, S1 = Shrubland 1, S2 = Shrubland 2, MS1 = Mountain Steppe I, MS2 = Mountain Steppe II,
W= Woodland, S= Scrub, A= Agricultural Area, R= Rock and Stones. Habitat classification based on Wallis de Vries et al. (1996). A value of log
w
i
> 0 represents positive selection, = 0 represents no selection, <0 represents avoidance. Squares indicate habitats were not selected at all, but were
arbitrarily assigned a value of 0.05 so that they appear on the figure, circles indicate w
i
values significantly different to zero (v
2
P< 0.05), triangles
indicate insignificant w
i
values or where expected values <5.
S.R.B. King, J. Gurnell / Biological Conservation 124 (2005) 277–290 287
4.3. Vegetation use
Plants change in nutrition and palatability through
the year (Novellie and Winkler, 1993), and in Hustai
National Park the takhi preferred some species in their
early development stages, and avoided others after they
had flowered (Salter and Hudson, 1979; Duncan, 1983;
Miller, 1983b; Putman et al., 1987; Enkhee, 1998). The
takhi selected vegetation classes dominated by grasses
and fescues which have a high fibre and crude protein
content (van Dierendonck and Wallis de Vries, 1993).
Although grazing was observed on all vegetation classes
overall, takhi did not select all vegetation classes in every
year, nor graze on all vegetation classes present.
Vegetation available to takhi affected their altitudinal
movements through the year. They spent more time at
low to mid elevations to shelter from inclement weather,
and to feed on the most nutritious vegetation that was
found in the Lowland steppe (Salter and Hudson,
1978). All the takhi had good condition throughout
the year, and in the spring tended to have retained more
body fat than local domestic horses (SRBK, unpubl.).
As the takhi population grows there is potential for
overgrazing to occur, especially on vegetation classes
in the valleys that are preferred. Thus management will
be needed in the future to ensure that this is kept under
control.
5. Conclusions and the future of takhi at Hustai National
Park
So far, the introduction of takhi to Hustai National
Park seems to have been successful, although this assess-
ment is based on a time period that only approximates
one generation (see Gross, 2000). Despite this, there is
little evidence that takhi have lost any ability to survive
in the wild through many generations in captivity. All
harems bar one were given a soft release, and main-
tained in acclimatisation enclosures for at least six
months, thus allowing animals to get used to both the
climate and disease vectors. The importance of a suffi-
cient acclimatisation period is shown by the greater suc-
cess of harems, such as PatronÕs, ParitetÕs and KhaanÕs,
which were acclimatised for two years, compared to the
those that had a hard release (TurgenÕs harem) or were
acclimatised for only six months (Mark and BohemianÕs
harems). These harems suffered greater mortality, with
only two horses from Turgen and BohemianÕs harem
surviving a year after release, and only one from MarkÕs.
Local diseases were the main cause of mortality, sug-
gesting knowledge of potential diseases should be ob-
tained before animals are brought to the release site,
and continuous monitoring post-release with immediate
veterinary attention is desirable. Although target release
sites should be surveyed for potential predators, in this
case predation was not a major cause of mortality. How-
ever this is not true for all introductions (e.g., Matson
et al., 2004).
The horses in this study tended to remain near their
acclimatisation enclosure upon release, although they
did move further away over time. Thus repeated use of
a release enclosure could lead to intraspecific conflict
and overexploitation of the nearby habitat. However,
this does not seem to have occurred at Hustai National
Park (enclosure 1 was used in 1994, 1995 and 1998, and
enclosure 2 in 1994 and 1998; enclosures 4 and 5 were
only used once); there was little overlap between ranges
of different harems, although exclusive range use did not
occur. All harem ranges at Hustai National Park con-
tained access to a water source, and this has also been
seen in feral horse studies. Releases should therefore
be near a permanent water source, and this could be
used to encourage animals to move further from a re-
lease enclosure. In addition shade areas and access to
minerals seemed necessary for each harem. In compari-
son with feral horse studies, it appears that range size
depends on habitat quality, and this should be taken
into account in assessing target release sites. Impor-
tantly, takhi appear to use their habitat in a similar
way to feral horses, so management techniques of feral
populations could be applied.
It has been suggested that Hustai National Park
(area 570 km
2
) should be able to support a population
of 400–500 horses (Bouman, 1998), equivalent to a
density of 0.9 horses km
2
or 42 harems with the cur-
rently observed 12 horses harem
1
. However, because
of the topography and availability of water, Hustai
National Park may not support 500 horses. For exam-
ple, in the western section of Hustai National Park
there are streams in most valleys, but in the east (c.
144 km
2
) there are only three ephemeral streams.
Horses require a reliable water source, along with
the presence of good quality vegetation and places
for resting both by rocks near ridges and among trees.
At present, the takhi population appears well below
carrying capacity. Taking the rather tentative 2002–
2004 estimate of per capita growth rate per year of
0.066, it will take 15.5 years for the population to
reach a carrying capacity of 450 horses.
A future problem is the likelihood of hybridisation
with domestic horses as the takhi expand beyond the
Park boundary. Local people value hybrid offspring.
Hybrids will need to be identified and either controlled,
neutered or prevented from re-joining the population.
Contraception may be a better way of limiting the pop-
ulation than removing animals (e.g., Gross, 2000). Fu-
ture studies at Hustai National Park will provide
insights into dispersal, harem formation, and the effects
of intraspecific competition as the population increases.
In addition studies of the horses over the winter may
provide more insights into the habitat use and any
288 S.R.B. King, J. Gurnell / Biological Conservation 124 (2005) 277–290
changes in social structure during this critical season.
Takhi born in Hustai National Park could be used for
future reintroductions in other parts of Mongolia as
they will already have immunity to babesiosis and will
have acquired the skills necessary to survive the varied
seasons.
Acknowledgements
We thank the Foundation Reserves Przewalski Horse
for their financial support and the opportunity of study-
ing the horses at Hustai National Park, as well as access
to past data. Thanks to the Mongolian Association for
the Conservation of Nature and the Environment and
the staff at Hustai National Park for all their help dur-
ing the study, and thanks especially to the Hustai Na-
tional Park biologists and rangers for collecting
invaluable data, to Tessa Roos, and to Marc Hooger-
werf for advice on using ArcView. Funding for this
study also came from IFAW, Queen Mary Expeditions
Committee and Marwell Preservation Trust.
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... In reintroduced and endangered species habitat use is one of the key studied factors [28,29]. In equids, forage abundance is the most important determinant for habitat use because of their digestive anatomy [30,31] and they may spend more than half of their daily time budgets grazing to get enough nutrients [32]. However, inconsistent and contradictory information has been described by previous observations of the feeding behaviour and daily budget of horses (feral horses [24][25][26][27][28][29][30][31][32][33][34]; wild horses [25,32,35]:). ...
... In equids, forage abundance is the most important determinant for habitat use because of their digestive anatomy [30,31] and they may spend more than half of their daily time budgets grazing to get enough nutrients [32]. However, inconsistent and contradictory information has been described by previous observations of the feeding behaviour and daily budget of horses (feral horses [24][25][26][27][28][29][30][31][32][33][34]; wild horses [25,32,35]:). In Hustai National Park, Mongolia, harems of Przewalski's horses usually feed in the morning and evening, before and after the walk to the water source [25,35]. ...
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Background Reintroduction is a common technique for re-establishing threatened species. However, the adaptation to novel habitats with distinct conditions poses a risk of failure. Weather conditions affect the behaviour of animals, and thus, their adaptation to new conditions and survival. Reintroduced Przewalski’s horses living in Mongolia’s continental arid climate with extreme temperature and precipitation variability, serve as an ideal model species for studying the behavioural response of selected groups to these harsh conditions. Methods The research was conducted in The Great Gobi B Strictly Protected Area, Mongolia. In summer 2018, three groups were recorded (Azaa, Tsetsen and Mares18) involving 29 individuals. In Spring 2019, 4 groups were recorded (Azaa, Tsetsen, Hustai1 and Mares19) involving 34 individuals. In Autumn 2019, 4 groups were recorded (Azaa, Tsetsen, Hustai2 and Tanan) involving 35 individuals. Thirteen weather variables were recorded in 10-min intervals, together with the percentage representation of selected behavioural categories (feeding, locomotion, resting, and social). The effect of weather on behaviour was analysed through GLMM. Influence of the group-history factors (recently reintroduced, long-term reintroduced and wild-born) was also analysed. Results Feeding significantly increased with cloudy and windy conditions and was more frequent in autumn than spring and summer. Locomotion was positively explained by temperature and cloudiness and was higher in summer than spring and autumn. Resting behaviour decreased with altitude and cloudiness, and the dispersion of the group was lower when resting. Increased social interactions were observed with higher temperatures and were more frequent in summer compared to spring and autumn. Differences were found in the display of the behaviours among the selected harems, showing interesting patterns when grouping them according to their origin and experience. Conclusions Weather patterns seem to influence the behaviour of Przewalski’s horse. These results might assist in further management plans for the species, especially in the view of intensifying climate change and alteration of weather patterns. As previously suggested, after approximately 1 year, horses adapt to novel conditions and display the typical behavioural pattern of wild-born Przewalski’s horses.
... Many studies reported the proportion of individuals remaining in the release area but do not consider the natural dispersal tendencies of the animals (e.g. Ostro et al., 2000;King and Gurnell, 2005;Hardman and Moro, 2006). It is often true that dispersal from a release site can be associated with a high mortality rate (Bright and Morris, 1994), but in highly territorial animals such as slow lorises, secondary dispersal appears to be a natural behavior. ...
... immediate) release since it reduces postrelease dispersal (e.g. King and Gurnell, 2005;Tuberville et al., 2005;Kenyon et al., 2014). Other studies, however, found no effect (e.g. ...
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Unmonitored release is a common practice, especially in small animals, that present a series of adverse conditions if not well-planned. Small research centers and non-governmental organizations in developing countries often receive animals that are then subject to unmonitored releases. We explored the patterns of post-release and natal dispersal in the Javan slow loris, a Critically Endangered venomous and territorial mammal that is highly threatened by wildlife trade. We then determined the importance of health status and human habituation for the survival of translocated and natally dispersing animals. We collected data from 2012 to 2018 on pre-release and pre-dispersal health conditions and human habituation, post-release and post-dispersal presence of wounds, behavior, and ranging patterns of 11 translocated and 11 natally dispersing individuals and compared them with 12 stable resident individuals. Translocated animals had a larger home range size (15.9 ± 4.1 ha) and higher wound presence during recaptures (0.47 ± 0.13) than stable resident individuals (3.2 ± 3.0 ha; 0.10 ± 0.06) but they did not differ from natally dispersing individuals (13.8 ± 3.7 ha; 0.28 ± 0.11). Both translocated and natally dispersing individuals can move to a different habitat type compared to their release area or natal range. The fate of both translocated and natally dispersing individuals was influenced by their health state (p < 0.001), and human habituation significantly affected the possibility of being captured for wildlife trade of translocated individuals (p = 0.048). We highlight the importance of considering natal dispersal, health state, and human habituation before the release of small animals to avoid death and capture for wildlife trade.
... Habitat selection and use are primarily influenced by forage availability and quality [91,92], with distance to water also being important [93]. The primary determinant of habitat use in free-roaming horses in all seasons has been shown to be availability of preferred forage [34,37,[94][95][96]. ...
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A detailed understanding of what is usual for a species under optimal conditions is critical for identifying and interpreting different features of body function that have known impacts on animal welfare and its assessment. When applying the Five Domains Model to assess animal welfare, the key starting point is therefore to acquire extensive species-specific knowledge relevant to each of the four physical/functional Domains of the Model. These Domains, 1 to 4, address areas where objective information is evaluated and collated. They are: (1) Nutrition; (2) Physical environment; (3) Health; and (4) Behavioural interactions. It is on the basis of this detailed knowledge that cautious inferences can then be made about welfare-relevant mental experiences animals may have, aligned with Domain 5, Mental State. However, this review is focused entirely on the first four Domains in order to provide a novel holistic framework to collate the multidisciplinary knowledge of horses required for undertaking comprehensive welfare assessments. Thus, inferring the potential mental experiences aligned with Domain 5, the final step in model-based welfare assessments, is not considered here. Finally, providing extensive information on free-roaming horses enables a better understanding of the impacts of human interventions on the welfare of horses in both free-roaming and domestic situations.
... Furthermore, we observed very low overlap of core areas, indicating exclusive use of core areas by bands of wild ponies in these areas as it was described for Exmoor ponies by Gates (1979), but contrary to what has been observed for other free roaming horses (Tyler 1972;Berger 1989;Linklater 2000). Previous studies reflect that free roaming horses in large open areas live in bands that largely or entirely overlap their HRs (Berger 1986, Linklater 2000Zabek 2015) and a similar pattern was observed for Przewalski horses (King and Gurnell 2005;Kerekes et al. 2021). However, exclusive use of HR and core areas has been described as an atypical spatial behaviour of dense populations of feral horses confined artificially (Linklater et al. 2000). ...
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Large herbivores are key regulators of open habitats across the world. Free roaming ponies have a prominent ecological role in many Atlantic landscapes, where different habitats with conservation interest are linked to ponies’ occurrence. The traditional management of wild ponies, which implies minimum human intervention, is declining in Galicia, NW Spain. Changes in the management regimes include the confinement of ponies in fenced areas, the use of improved pastures (IPs) and rotation between fields. Indirect effects of these changes are expected on the ecological condition of important habitats for conservation such as dry and wet heathlands and bogs. We studied social structure, spatial ecology and habitat use in 29 mares fitted with global positioning system (GPS) collars and field observations in two areas of Galicia dominated by wet heaths and blanket bogs (Xistral), and dry heaths (Sabucedo). We used spatial location and field observations to identify each band, and calculated band size, sex ratio, home range (HR) and core areas size and overlap, and habitat use. We addressed differences and adjusted Generalized Linear Models (GLMs) for these variables as functions of the type of management: free roaming vs rotation, use of IPs, fencing, and available ranging area. Larger bands were found in smaller commons, fenced and with rotation management. Home ranges, but not core area, varied as a function of the available ranging area. Bands overlap more on fenced areas with rotation management. Increasing management may concentrate grazing pressure by reducing HR and increasing bands overlapping areas, and this may have a long-term effect on habitat quality and conservation.
... All Przewalski's horses alive today are descendants from only 13 individuals that were the nucleus for captive breeding [1,3,6,8,9]. After another 20 years of captive breeding on four different continents, including Asia, the total number of horses rose to almost 1000 individuals [3,6,7,[10][11][12][13][14], and many organizations have since then also attempted to release the horses in semiwild habitats such as the Kalamaili Nature Reserve in Xinjiang, China. Although this may be a step in the right direction, these populations remain somewhat dependent on human support for their survival. ...
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Recent technological innovations have led to an upsurge in the availability of unmanned aerial vehicles (also known as drones and hereafter referred to as UAVs)—aircraft remotely operated from the ground—which are increasingly popular tools for ecological research, and the question of this study concerns the extent to which wildlife responses might allow aerial wildlife monitoring (AWM) by UAVs. Our experiment tests the hypothesis that the wildlife-UAVs interaction depends strongly on flight altitude that there may be a lowest altitude range for which the ungulates are not exceedingly disturbed, dictating a practically achievable level of discernibility in flight observation, for this question might influence the future viability of the UAVs in the study and protection of the other wildlife in China’s semiarid ecosystem. We examined the behavioral responses of a group of enclosed Przewalski’s horses (Equus ferus przewalskii) to the presence of different in-flight UAVs models by conducting flights at altitudes ranging from 1 to 52 meters and recorded the heights at which each horse reacted to (noticed and fled) the UAVs. All horses exhibited a stress response to UAVs flights as evidenced by running away. The results suggest strong correlations between flight altitude and response across the different subjects that adults generally noticed the UAVs at the larger heights (20.58 ± 10.46 m) than the immature (4.67 ± 0.87 m). Meanwhile, reaction heights of females (15.85 ± 6.01 m) are smaller than that of males (26.85 ± 18.52 m). Supported by their biological roles in herds (e.g., males must give protection to his entire herd while females are purely responsible for their offspring), our results also show that age, closely followed by gender, are the two most significant elements that determine a horse’s level of alertness to the UAVs. This research will help future scientists to better gauge the appropriate height to use a drone for animal observation in order to minimize disturbance and best preserve their natural behavior. 1. Introduction 1.1. Przewalski’s Horses Przewalski’s horses (Equus ferus przewalskii), also known as Mongolian wild horses or Dzungarian Horses, are the last surviving subspecies of wild horses. Although initially sighted in the 15th century, these horses were not scientifically documented until 1880, and these horses were sighted in the Mongolian Gobi Desert, by a Russian officer, Colonel Nikolai Przhevalsky, so the horses were named after him [1, 2]. After being captured and transferred to Europe in the early 1900s, Przewalski’s horses soon became endangered due to habitat loss, over hunting, and competition with livestock [1–3]. The last recorded sighting of a wild Przewalski’s horse occurred in the Dzungarian Gobi of Mongolia in 1969 [4], and since then, this species has been extinct in the wild with only a few remnant populations existing in small captive breeding herds in Western countries, making the survival of the taxon possible [5–7]. All Przewalski’s horses alive today are descendants from only 13 individuals that were the nucleus for captive breeding [1, 3, 6, 8, 9]. After another 20 years of captive breeding on four different continents, including Asia, the total number of horses rose to almost 1000 individuals [3, 6, 7, 10–14], and many organizations have since then also attempted to release the horses in semiwild habitats such as the Kalamaili Nature Reserve in Xinjiang, China. Although this may be a step in the right direction, these populations remain somewhat dependent on human support for their survival. Therefore, recommendation for the success of this project states to limit the number of domestic animals in the area, in order to ensure a long-term self-sustaining population of Przewalski’s horses [1–3]. 1.2. Previous Use of Unmanned Aerial Vehicles in Studies of Wildlife Research Unmanned aerial vehicles (UAVs) have the potential to revolutionize the way in which research is conducted in many scientific fields [15, 16]. UAVs have proven to be useful devices for the observation of wildlife, in particular the production of systematic data with high spatial and temporal resolution because the devices can access remote or difficult terrain [17], collect large amounts of data for lower cost than traditional aerial methods, and facilitate observations of species that are wary of human presence [18]. Currently, despite large regulatory hurdles, UAVs are being deployed by researchers and conservationists to monitor threats to biodiversity, collect frequent aerial imagery, estimate population abundance, and deter poaching [19–24], but with the widespread increase in UAV flights, it is critical to understand whether UAVs act as stressors to wildlife and to quantify that impact. It is likely that UAVs could also have unwanted and unanticipated risks on wildlife and their delicate ecosystems. Research has shown that retaliation against UAVs by terrestrial mammals was different from that of marine mammals and aquatic birds [25, 26]. For example, a study into the free-roaming American black bears (Ursus americanus) proved that the presence of UAV flights brings significant distress to the physiological state, which often does not manifests itself in terms of behavioral changes, proving difficult for observers to discern [27]. Furthermore, a study on guanacos (Lama guanicoe) revealed that low-flying UAVs at any speed, as well as high-flying UAVs at an accelerated velocity, caused a disturbance to the guanacos’ behavior [28]. Although UAVs have potential for success, a great deal of uncertainty still surrounds their use, and scientists must be cautious of the impact that they bring to each new investigation, especially when endangered species or ecologically sensitive habitats are involved [27]. Moreover, it is vital that UAVs are manufactured and selected to minimize visual and audio stimuli in order to reduce disturbance of wildlife. Shape, volume, and color are all factors to be taken into consideration thus to mimic nonthreatening wildlife native to the studied habitat in order to decrease disruption [29]. Sporadic movements and threatening or alarming trajectories should be avoided at all costs, and if an operation proves to be excessively disruptive, it must be ceased immediately [27]. With these basic regulations in place to ensure safety and ethicality, UAV flight are sure to make steady progress and evolve into more effective devices for the study of animal behavior. 1.3. Summary and Hypothesis As the last surviving subspecies of wild horses, Przewalski’s horses are a great subject of interest to the scientific community. Despite extensive studies especially on their social behavior and time budget [30–38], Przewalski’s horses remain somewhat of a mystery to scientific world today, primarily because of their rarity. The purpose of our study was to better understand Przewalski’s horses, in particular their response to UAVs under semireserve conditions. Previous studies of Przewalski’s horses and domestic and feral horses (Equus caballus) have shown left side bias in vigilance responses with horses reacting sooner when approached from the left side by novel stimuli [39–41]. In this study, we wanted to look at the influence of the height of novel stimuli. We specifically focused on discerning how individual horses reacted to the height of UAVs and which factors influenced their reactions, whether it be age, gender, or the herd that they resided in. Our hypothesis is that wildlife-UAVs interaction depends strongly on flight altitude, that flying too low could excessively disturb them and that there may be a lowest altitude range for which the ungulates are not exceedingly disturbed. Additionally, we hoped to learn the height at which UAV flights could operate without disturbing the horses and whether UAVs can be a reliable tool in the observation of their behavior and other wild wildlife. We also hypothesized that horses would respond to the UAV flight in one of four ways: no discernable responses (positive) and discernable responses (negative) (i.e., attention, eye/head movement and/or turning of the upper body, increased movement rates, and/or moving away slowly from the UAVs). We also hypothesized that the degree of responses would vary with respect to age and gender. 2. Materials and Methods The design of the experiment began determining a range of flight altitudes. The range of flight altitudes was determined through preliminary tests, flying at altitudes from 1 meter to 100 meters in 10 m increments. We determined that 1–5 m was a suitable lower bound because of safety concerns and that 50–60 m was a suitable upper bound because negative response had already significantly dropped off by that altitude. 2.1. Study Area This study was conducted at three separate enclosures of the Wild Horse Breeding Research Center of Xinjiang Uygur Autonomous Region, China (44°12′20.5′N, 88°44′52.8′E). This area has a semiarid, desert climate with great seasonal temperature differences: the hottest recorded summer temperatures 40°C, 104°F, and winter temperatures often fall below −15°C. To maintain natural breeding patterns and social structures of Przewalski’s horses, the horses typically remained in their native herd following birth to be raised by their mothers. Once they reached a mature age, they would be reallocated to herds corresponding to their genders. Occasionally, mares would be regrouped to create a novel mixed herd, while stallions would be reintroduced to such mixed herds to determine dominance. The defeated stallion returns to his single-sex herd, while the victorious stallion gains control over his new mixed-sex herd. 2.2. Data Collection The unmanned aerial vehicle (UAVs) we used was a Mavic 2 Zoom drone, powered by a 1/2.3 inch 12 MP sensor with up to 4x zoom, including a 2x optical zoom (24–48 mm) to capture all shorts from wide angle to midrange. The 4x lossless zoom and 2x optical zoom enabled a closer, high-definition view of faraway subjects while maintaining our distance to decrease the disruption to our horses. The Mavic 2 cameras utilized DJI’s latest 3-axis gimbal technology and recorded videos at higher bitrates with advanced H.265 compression. Videos in H.265/HEVC codec maintain 50% more information than videos in H.264/AVC, which leads to better preserved details allowing us to clearly recognize behavioral signals in our horses. Additionally, FOC sinusoidal drive ESCs and low-noise propellers reduce flight noise, thus reducing disturbance to the horses. Binoculars (10x magnification) were used when needed. Horses were followed on foot and watched at distances of 30–100 m. Individual horses were identified using sex, size, color, and distinguishing markings. UAV flights were conducted from August 1, 2020, to December 08, 2020, at consistent times during daylight hours in order to reduce the influence of other factors that may affect the reaction times of Przewalski’s horses. The research team consists of four personnels: (1) a pilot, (2) a primary observer, (3) a ground camera operator, and (4) a data recorder. The pilot was responsible for flying the UAVs. Only one horse was scored when the drone was flown over a herd. Flights were made of 62 horses including 53 adults and nine immature horses (adult male, N = 9; immature male, N = 5; adult female, N = 44; and immature male, N = 4) living under natural social conditions at three separate enclosures. The ages of the horses were obtained from records of the reserve association (horses ranged from 1 day to 2 years were categorized as immature until they had left their natal band). Since these horses have been studied extensively, they were habituated to observation and were not highly vigilant and reactive to the presence of the observer, who could watch from a distance up to as little as 20–50 m while remaining stationary. The UAVs were launched approximately 100 m from the targeted location of the horse. The total dataset consisted of 62 flights that ranged in altitude from 1 to 100 meters. Each flight began with taking off from a launch point, flying about 100 m high from a focal individual animal, changing altitude as appropriate when passing over the subject. The UAVs were flown over the animal or herd at 10 m/s. The next pass was flown at another randomly selected altitude. If an affirmative response was invoked, the drone would wait at a distance, while the animals would settle back to a sedentary behavior. We used the live video feed which also offers a recorded video after a completed flight and the location stamp, which informs us of the exact height and angle of the drone to determine the vertical distance from our subjects, as well as the exact measurements at which they individually displayed the two levels of reaction that we were measuring. To avoid confusion, the term “response” is used here for all levels of behavioral responses, as it is for a broad range of species to refer to an animal interrupting feeding, by lifting or cocking its head, and to attend to its surroundings [41]. Level 1, known as the alert height, was determined through the eye and ear movements of Przewalski’s horses; these reactions can range from subtle ear swivels indicating auditory influence to indirect glances at the UAVs. Once a subject displays any of these reactions, it becomes aware of the existence of the drone, and the height of the UAVs was recorded as the alert height. Level 2, known as the run away height, was the height of the UAVs at which the subject physically displaced itself to avoid the UAVs. 2.3. Data Analysis Data were tested for normality with the one sample Kolmogorov–Smirnov test. Because all of the data showed a normal distribution, we used the t-test to detect the differences of alert height and flight initiation height between the age and sex. We then used the general linear model to test the effect of age, sex, and their interaction on alert height and flight initiation height. We accepted statistical significance at the level of , and all the data were analyzed using the SPSS 19.0 statistical package. 3. Results Przewalski’s horses responded to UAV flights in all 62 flights. Tables 1 and 2 display the comparative makeup of each response for the different altitudes. First, notice that the positive responses (no discernable movement and alert) increases with altitude, while the negative responses (discernable movement and alert) decrease with altitude. The results help us illustrate the general pattern of less negative response at higher altitudes supporting the hypothesis that altitude plays a large role in animal response from UAVs. Next, notice that the degree of response varied with respect to age and gender: the alert height (height for horse to notice UAVs) and run away height (height for horse to move away) of adult horses ranged 11–52 and 1–36 meters while that of immature ranged 3–6 and 1–2 meters, respectively. The result also showed that immature compared to adults showed a greater effect in males than in females (Table 1). Additionally, alert height and run away height for female of all tested herds ranged 8–30 and 1–4 meters, while for male ranged 29–52 and 4–36 meters, respectively, showing that male horses are more alert than the females (Table 2). In summary, age, sex, and their interaction have significant effects on alert height and run away height (Table 3). Variables Number Alert height (m) ± SE t-test Run away height (m) ± SE t-test Males Immature males 5 5.0 ± 0.71 1.40 ± 0.55 Adults males 9 40.33 ± 9.38 18.67 ± 12.80 Females Immature females 4 4.50 ± 0.50 1.25 ± 0.50 Adults females 44 16.68 ± 5.26 2.16 ± 0.83 Alert height, height for horse to notice UAVs; run away height, height for horse to move away (means with standard errors). “Number” refers to the number of flights: only one horse was scored when the drone was flown over a herd.
... Although horses are highly mobile, they tend to remain within home ranges (King 2002, King andGurnell 2005). Understanding space use is important to detect areas of high horse density, to assist managers in understanding how horses are using habitat, and to provide information about grouping of individuals. ...
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Feral horses (Equus ferus caballus) in the western United States are managed by the Bureau of Land Management (BLM) and United States Forest Service in designated areas on public lands with a goal of maintaining populations in balance with multiple uses of the landscape. Small, isolated populations can be at risk of extirpation from stochastic events and deleterious genetic effects resulting from inbreeding and reduced heterozygosity. The genetic diversity of feral horse herds is periodically monitored using blood or hair samples collected during management gathers (i.e., occasions when the herd is rounded up). We conducted a study to examine genetic characteristics of the feral horse population at the BLM Little Book Cliffs Herd Management Area (HMA) in Colorado, USA, using non‐invasively collected fecal samples. Additionally, we explored whether genotypes could be used to document space use and potential sub‐population development. We used a random sampling scheme, walking transects in sampling areas covering most of the HMA to find and collect fecal samples of all ages, except those that were deteriorating. We collected >1,800 fecal samples from across the study area in May, August, and October 2014. We then identified unique individuals using a suite of microsatellite loci. Our estimates of genetic diversity from fecal samples were higher than those reported from blood and hair samples taken during recent horse gathers, likely because our sample size and spatial distribution was larger. Genotypes revealed that some individuals were found only in certain parts of the study area and at a higher proportion than random; thus, they could be considered residents in those sampling areas. Using discriminant function analyses, we detected 5 genetic groups in the sample population, but these did not correspond to individuals in specific parts of the study area. Our results support the use of fecal DNA to augment direct observations of horse presence and could be used to detect habitat use and areas of high density. Non‐invasive techniques such as fecal DNA sampling can help managers decide whether new individuals need to be translocated to a closed population to maintain genetic diversity without the human safety and animal welfare concerns associated with gathers and invasive techniques. © 2021 The Wildlife Society. This article is a U.S. Government work and is in the public domain in the USA. We found higher estimates of genetic diversity from feral horse fecal samples collected non‐invasively than those reported from blood or hair samples collected during a gather; genotypes derived from fecal samples can be used to show feral horse space use. Fecal samples can therefore be used to examine genetic characteristics of a horse herd in the absence of a gather and can elucidate areas of high horse use.
... However, the habitat use by grazers within an enclosure is usually not homogenous (Gander et al., 2003;King & Gurnell, 2005). For understanding the influence of space use patterns of grazers on birds within a given grazed area, it may help to obtain position data of individual grazers. ...
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Grazing by large herbivores is increasingly used as a management tool in European nature reserves. The aim is usually to support an open but heterogeneous habitat and its corresponding plant and animal communities. Previous studies showed that birds may profit from grazing but that the effect varies among bird species. Such studies often compared bird counts among grazed areas with different stocking rates of herbivores. Here, we investigated how space use of Konik horses and Highland cattle is related to bird counts in a recently restored conservation area with a year-round natural grazing management. We equipped five horses and five cattle with GPS collars and correlated the density of their GPS positions on the grazed area with the density of bird observations from winter through the breeding season. We found that in the songbirds of our study site, both the overall density of bird individuals and the number of species increased with increasing density of GPS positions of grazers. Correlations of bird density with horse density were similar to correlations with cattle density. Of the eight most common songbird species observed in our study area, the Eurasian Skylark and the Common Starling had the clearest positive correlations with grazer density, while the Blackbird showed a negative correlation. Skylarks and Starlings in our study area thus seem to profit from year-round natural grazing by a mixed group of horses and cattle.
... En prenant en compte les exigences environnementales de ces deux espèces, nous pensons que leur chasse avait lieu pendant la belle saison, entre la fin du printemps et le début de l'automne. Nous n'avons pas pour l'instant de données sur l'âge de mort des animaux, qui pourraient alimenter cette thèse, car les analyses cémentochronologiques n'ont pas été réalisées à cause de la mauvaise préservation des ossements ; mais des études récentes sur les migrations saisonnières des chevaux sauvages et surtout des cerfs vont dans ce sens(King et Gurnell 2005, Luccarini et al. 2006, Kaczensky et Huber 2010. Le site est un emplacement stratégique du point de vue cynégétique, ce qui explique le choix des groupes qui y ont séjourné : lié à la chasse du cheval, Montlleó se situe sur sa route migratoire saisonnière entre la plaine de la Cerdagne et les piémonts pyrénéens.Les occupations humaines sur le site de plein air du Paléolithique supérieur ... La conquête de la montagne : des premières occupations humaines à l'anthropisation du milieuFig. ...
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CITATIONS 0 READS 5 8 authors, including: Some of the authors of this publication are also working on these related projects: Cultural behaviour of hunter-gatherer societies during the Early Upper Palaeolithic in northeast Iberia. Techno-morphological study of personal ornaments View project The importance of small prey in Iberian Palaeolithic hunter-gatherers sites: The bird assemblages from Arbreda Cave (Serinyà, Girona) View project
... En prenant en compte les exigences environnementales de ces deux espèces, nous pensons que leur chasse avait lieu pendant la belle saison, entre la fin du printemps et le début de l'automne. Nous n'avons pas pour l'instant de données sur l'âge de mort des animaux, qui pourraient alimenter cette thèse, car les analyses cémentochronologiques n'ont pas été réalisées à cause de la mauvaise préservation des ossements ; mais des études récentes sur les migrations saisonnières des chevaux sauvages et surtout des cerfs vont dans ce sens(King et Gurnell 2005, Luccarini et al. 2006, Kaczensky et Huber 2010. Le site est un emplacement stratégique du point de vue cynégétique, ce qui explique le choix des groupes qui y ont séjourné : lié à la chasse du cheval, Montlleó se situe sur sa route migratoire saisonnière entre la plaine de la Cerdagne et les piémonts pyrénéens.Les occupations humaines sur le site de plein air du Paléolithique supérieur ... La conquête de la montagne : des premières occupations humaines à l'anthropisation du milieuFig. ...
... En prenant en compte les exigences environnementales de ces deux espèces, nous pensons que leur chasse avait lieu pendant la belle saison, entre la fin du printemps et le début de l'automne. Nous n'avons pas pour l'instant de données sur l'âge de mort des animaux, qui pourraient alimenter cette thèse, car les analyses cémentochronologiques n'ont pas été réalisées à cause de la mauvaise préservation des ossements ; mais des études récentes sur les migrations saisonnières des chevaux sauvages et surtout des cerfs vont dans ce sens(King et Gurnell 2005, Luccarini et al. 2006, Kaczensky et Huber 2010. Le site est un emplacement stratégique du point de vue cynégétique, ce qui explique le choix des groupes qui y ont séjourné : lié à la chasse du cheval, Montlleó se situe sur sa route migratoire saisonnière entre la plaine de la Cerdagne et les piémonts pyrénéens.Les occupations humaines sur le site de plein air du Paléolithique supérieur ... La conquête de la montagne : des premières occupations humaines à l'anthropisation du milieuFig. ...
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The open air site of Montlleó (Prats i Sansor, Lérida, Spain) was discovered in 1998 and has been excavated since 2000 by a multidisciplinary team from the SERP research group, University of Barcelona. The presence of lithic and bone industries, as well as shell ornaments, have shown the importance of the site for hunter-gatherer Magdalenian groups and the role of the high Segre valley as a crossroad for lithic raw materials exchanges on the Segre-Têt axis through the Cerdanya valley. New radiometric dates (which situate the first occupations up to 22,000 cal BP) and the presence of new typological tools such as raclettes and Solutrean shouldered points lead us to think that human occupations were more ancient than first expected, going back up to the beginning of the Last Glacial maximum. The dispersion analyzes actually show the existence of two archeological levels separated by a short sterile hiatus. These data thus show that this site represents the oldest occupation of a mountain area during the LGM in the southeastern Pyrenean region.
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