B I O L O G I C A L C O N S E RVAT I O N 1 4 0 ( 2 0 0 7 ) 1 –7

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Effectiveness of population recovery projects based

on captive breeding

Paweł Adamskia,*, Zbigniew J. Witkowskib

a
Institute of Nature Conservation, Polish Academy of Sciences, al Mickiewicza 33, 32-120 Kraków, Poland
b
Faculty of Ecology and Environmental Management, Academy of Physical Education, al. Jana Pawla II 78, 31-571 Kraków, Poland

A R T I C L E I N F O A B S T R A C T

Article history: A recovery programme for a population of Apollo butterflies in the Pieniny National
Received 22 May 2006 Park was monitored over 12 years in order to assess its effectiveness. The programme
Received in revised form was based on population ecology and metapopulation theory, and its object was to
14 May 2007 replenish wild populations with captive-reared individuals. Thanks to the adopted mea-
Accepted 22 June 2007 sures, the local population initially numbering only 30–50 individuals increased to ca.
Available online 14 September 2007 1000 imagines, forming a functional metapopulation. Nevertheless, analysis of popula-
tion dynamics parameters indicated that only some of the subpopulations were stable;
Keywords: a significant percentage of the subpopulation fractions represented sink populations
Apollo butterfly whose survival depended either on an external supply of captive-reared individuals or
Population recovery on individuals from other source subpopulations. In some cases, intensively supplying
Metapopulation the subpopulation with captive-reared individuals resulted in a decrease of that deme’s
Captive breeding abundance.
Reproduction rate The results suggest that supplying captive-reared individuals to an endangered pop-
ulation may entail some risks and should be done with caution.
Ó 2007 Elsevier Ltd. All rights reserved.

1. Introduction
One of the most spectacular active measures employed to
protect an endangered species or population is to aid its
recovery with the use of captive breeding (Hutching, 1997;
Saint-Jalme, 2002). Projects of this kind require major finan-
cial outlays for breeding centre upkeep (Kleiman et al., 1991)
and long-term monitoring (Ramoto et al., 1993; Johnson,
1994). For these reasons, Caughley and Gunn (1996) suggest
that recovery projects should be rather short-term, serving
the immediate goal of taking a population out of extreme
danger. To optimize the financial resources spent on recov-
ery projects, it seems important to set clear-cut goals, which
upon being reached trigger plans to discontinue or suspend
extremely labour-intensive and costly measures (Machado,
1997). The plan should also indicate whether a given popula-
tion will be self-sustaining or will continue to rely on a sup-
ply of captive-bred individuals. This paper evaluates the
effectiveness of a 12-year project to restore the Apollo but-
terfly (Parnassius apollo) population in the Pieniny National
Park.
2. Study site
The Pieniny Mts. are a narrow limestone mountain range
straddling the Polish-Slovakian border (Fig. 1). This area is
inhabited by one of the best-known Apollo butterfly popula-
tions in the northern part of the Carpathians (Nowicki-Siła,
_
1865, 1870; Sitowski, 1922; Zukowski, 1959; Witkowski, 1986;
Adamski and Witkowski, 1999a). The population is made up

* Corresponding author: Tel.: +48 126321101; fax: +48 1266322432.

E-mail address: adamski@iop.krakow.pl (P. Adamski).
0006-3207/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2007.06.027
2 B I O L O G I C A L C O N S E RVAT I O N
Fig. 1 – Location and structure of the Pieniny Apollo butterfy
metapopulation 1 – subpopulations, 2 – functional
population centers: W –Western, C – Central, E – Eastern, G –
‘‘Miedzy Grabczychami’’, ? – possible, but unverified site; 3 –
Border Pieniny National Park.
of the subspecies Parnassius apollo frankenbergeri (Slaby, 1955),
which inhabits only Pieniny National Park and the Haligovske
Hory mountain massif in Slovakia. Population decline and
habitat shrinkage were observed as early as the beginning
of the 20th century. These processes quickened significantly
in the 1950s. Significant changes in land use exacerbated
the Apollo’s decline. These changes included abandonment
of grasslands previously used as meadows or pastures and,
in the 1960s, afforestation of xerothermic grassland and stony
debris, critical habitats for the Pieniny Apollo (Witkowski and
Adamski, 1996; Witkowski et al., 1997). The Pieniny Apollo has
also been under strong pressure from butterfly collectors
(Adamski and Witkowski, 1999a).
3. Study population
In the late 1980s and early 1990s, the Apollo butterfly popula-
tion in the Pieniny National Park was near extinction. The to-
tal area of habitats optimal for the Apollo butterfly was
estimated at about 10 ha. The majority of the sites were frag-
mented and transformed. The whole population inhabited
only one site in the Trzy Korony massif (eastern part of meta-
population; Fig. 1), and its abundance was estimated to be be-
tween 20 and 30 imagines (Witkowski et al., 1997). Apart from
the dramatic drop in population numbers, there were some
other disquieting phenomena: a large proportion of individu-
als with pathologically developed wings in the population
(Adamski and Witkowski, 1999b), and an increase in the aver-
age fluctuating asymmetry of individuals (Adamski and Wit-
kowski, 2002). Those occurrences were probably linked with
genetic erosion, but there was no direct analysis of that
population.
In order to prevent the extinction of the population, a
recovery programme was launched in 1991. For the proper
functioning of the restored Apollo butterfly metapopulation
in the Pieniny National Park, four conditions would have to
be met:
1 4 0 ( 2 0 0 7 ) 1 –7
1. Restoration of subpopulations of the species at most of the
critical sites that meet its habitat requirements (Witkowski
et al., 1997).
2. Augmentation of the whole metapopulation to a level
close to the carrying capacity of the Pieniny National Park,
estimated in a pilot project to be 1000–1500 imagines (Wit-
kowski et al., 1992).
3. Restoration of a functioning metapopulation in which
exchange of individuals between subpopulations occurs
(Witkowski et al., 1997).
4. Creation of at least one large major site exhibiting features
of a metapopulation source within the Pieniny metapopu-
lation (Pulliam, 1988).
The degree to which these conditions have been met can
serve as a measure of the effectiveness of the programme.
Two principal measures were employed to achieve them (Wit-
kowski and Adamski, 1996):
Reconstruction of xerothermic grassland the crusial hab-
itat for the apollo habitat of the Apollo butterfly. Two factors
basic to the presence of the Apollo were analysed: the abun-
dance of the Apollo’s host plant, Sedum maximum, and the
extent of xerothermic grassland. Abundance and distribu-
tion analyses showed that the food plant is quite common
but that much of it is unavailable to the Apollo because it
grows in areas afforested by natural succession or by man-
agement activities (Witkowski et al., 1992). In such cases,
conservation activity concentrated mainly on removing
trees and shrubs from the historical localities of the Apollo
butterfly (Witkowski et al., 1997). By 2003 the area of the
Apollo’s habitat increased by ca. 19 ha. Moreover, it con-
sisted of a more compact set of sites, and stepping-stones
ßt between those sites had been established.
Introducing captive-bred individuals into the habitat. The
natural population was supplemented with captive-bred
individuals from 1992 till 2001. The captive breeding pro-
gramme was established in 1991 based on individuals taken
from the last surviving Apollo deme from the ‘‘E’’ part of the
population. Until 1995, replenishment was restricted to two
areas – ‘‘E’’ and ‘‘G’’. In 1996 the first captive-reared individ-
uals were introduced to restored habitats in the ‘‘W’’ and
‘‘C’’ parts of the metapopulation. Supplementation of the
‘‘W’’ and ‘‘G’’ parts of the metapopulation ceased in 1997
(Adamski and Witkowski, 1999a) (Fig. 2). In 1995, in order
to obviate the potential effects of genetic erosion, it was
decided to include in the breeding programme some
Fig. 2 – Intensity of captive- reared support of the wild
population.
 B I O L O G I C A L C O N S E RVAT I O N
individuals from the nearest population of the same subspe-
cies, obtained from Haligowskich Skaly (Slovakia). After this
measure, in 1996 three breeding lines were obtained: Polish,
Slovak and mixed (Witkowski and Adamski, 1996; Adamski
and Witkowski, 1999a).
In the beginning of the project the idea of artificially
increasing food plant abundance was considered. However,
abundance studies of the stonecrop (Sedum maximum) (Wit-
kowski et al., 1992) indicated that this perennial plant was
still quite abundant at the sites, including highly altered ones.
These activities were coupled with studies aimed at mon-
itoring the effect of the activities and solving problems
appearing during their implementation. After 12 years of
intensive work, the data gathered to date warrant an assess-
ment of whether the restored metapopulation is stable and
can survive without permanent support from captive-reared
individuals.
The Pieniny metapopulation, restored as a result of prepa-
ratory activities and introduction, can be divided in three
large functional population patches (Fig. 2).
The eastern part of the metapopulation (E), inhabiting the
Trzy Korony massif and Sobczanski gorge. The area has some
nine screes of various sizes accompanied by xerothermic
grassland.
The central part (C), including the xerothermic grassland-
covered slopes of Gorczynski gorge and similar habitats of
Nowa Góra, Macelowa Góra and the Cyrlowa Skałka hills.
The western assemblage (W) of demes on fairly extensive
and compact screes located on the Ubszar, Długa Grapa, Flaki
and Cisowiec hills, and the slopes around Czorsztyn castle.
The restored metapopulation occupies yet another single
site on a large a large area of stonedebris called Miedzy Grab-
czychami (G); despite it’s proximity to the Trzy Korony massif
(E) (Fig. 1), it is evidently isolated from it by the relief of the
area. Currently the area suitable for the Apollo butterfly cov-
ers about 40 ha, under the management of National Park
staff.
4. Methods
The population numbers were determined by marking and
recapturing imagines (Witkowski et al., 1997). During the per-
iod when the imagines were present, each of the potential
sites of Apollo butterflies were visited twice a week for 1–
3 h, depending on site size.
During these visits, all observed Apollo butterflies were
captured and the following data were registered for each
catch:
(1) The date and time of capture.
(2) The code of the individual. At capture each individual
was marked with a unique code allowing its unambig-
uous identification. The individuals obtained from
breeding in captivity were marked before their release
to sites.
(3) The sex of the individual, and in females the presence
or absence of the sphragis, a structure formed by the
male over the femaletßs abdomen to prevent her from
remating and displacing his sperm.
1 4 0 ( 2 0 0 7 ) 1 –7 3
(4) The site of capture, and for larger sites also the approx-
imate location within the site. The site where capture
occurred, and in larger sites also the approximate loca-
tion within the site.
The collected data were used to estimate the population
numbers of the Pieniny metapopulation through the Jolly
method, using the Jolly software package (Pollock et al.,
1990). This method, based on the ‘‘open population’’ was
checked because of significant migration rate within the pop-
ulation and the emergence of newly hatched individuals dur-
ing the sampling period; for this reason, method B was used.
The total population abundance in the whole sampling period
was extrapolated from the estimated population abundance
in a given sampling (Seber, 1982; Pollock et al., 1990). The
marking of individuals also revealed cases of migration be-
tween sites.
To determine the stability of the restored population, a
modified net reproduction rate coefficient (Krebs, 1994) was
calculated, taking into account the presence of captive-bred
individuals on the site
Wtþ1
Et ¼ ð1Þ
Wt þ Ct
where Et represents the yearly effectiveness of restoration
activity, Wt the number of wild individuals in the population
in year t, Wt+1 the abundance of the wild population in the
next year, and Ct the number of captive-reared individuals re-
leased to the site in year t.
The coefficient was calculated for the whole population as
well as for each of the four parts distinguished within it.
In each year, a migration coefficient was determined for
the metapopulation, indicating the fraction of marked indi-
viduals that undertook migration outside their original habi-
tat patch. The coefficient was calculated separately for
short-distance (within metapopulation centres) and long-dis-
tance migration (between these centres).
To assess the impact of introducing captive-reared individ-
uals into the population, regression analysis was performed
to determine the effects of wild population size and introduc-
tion intensity on reproduction rate in a given part of the
metapopulation, as well as on population number in the fol-
lowing year.
5. Results
During the period in which the reintroduction measures were
carried out, the range of the Pieniny metapopulation of the
Apollo butterfly expanded from the single subpopulation in
the Trzy Korony massif to more than a dozen subpopulations
living at almost all the major sites potentially suitable for this
species within the Pieniny National Park (Fig. 3).
The estimated total number of the Pieniny metapopulation
increased from a mere 30 individuals in 1991 to over 1200 in
2002 (Fig. 4a), with the greatest increase occurring in the Trzy
Korony massif, where ca. 50% of the metapopulation live
(Fig. 4b).
The values for short-distance dispersion within metapop-
ulation centres calculated since 1996 have fluctuated between
30% and 40% of the individuals found (Table 1). The
4 B I O L O G I C A L C O N S E RVAT I O N
Fig. 3 – Expansion of the metapopulation range during the
recovery process.
Fig. 4 – Changes in population abundance during the
reintroduction process: a — whole metapopulation 1 –
number of marked individuals, 2 – estimated population
size, 3 – +/- standard error of the estimated population size,
b — particular metapopulation centers.
1 4 0 ( 2 0 0 7 ) 1 –7
long-distance migration coefficient increased initially, reach-
ing a maximum 4.7% in 1999, and then declined to 2% in
2002, but the ratios of long-distance migrants between partic-
ular non-zero years did not differ significantly (v2 = 9.267,
df = 8, p = 0.3206) (Table 1). In 1996, all individuals undertaking
migration were those reared in captivity from either the Slo-
vak or the mixed line. The sex ratio of the migrants did not
differ from those of all marked specimens (v2 = 226, df = 1,
p = 0.1448).
The value of the reintroduction effectiveness index for the
whole metapopulation exceeded 1 only in 1992 and 1997. The
values of these indices for particular metapopulation centres
are given in Table 2.
Regression analysis indicates that the number of wild indi-
viduals emerging at a site in a given year was closely related
to the number of a given subpopulation in the previous year
(N = 48, betawild = 0.431, pwild < 0.0001 SEbetawild = 0.042),
whereas the effect of captive-bred individuals introduced to
the site in the previous year was not significant (N = 48,
betacaptive = 0.084, pcaptive = 0.1366 SEbetacaptive = 0.055).
Since population abundance at the sites often distinctly dif-
fered between years, the data were divided into two groups by
the median of the number of individuals observed at the given
site. When the number of observed individuals was lower than
the median calculated for the site, population abundance in
that year was positively related to the same value in the previ-
ous year (N = 29, betawild = 0.790, pwild < 0.0001, SEbetawild =
0.117, betacaptive = 0.115, pcaptive = 0.0228, SEbetacaptive = 0.047).
When the population number at the site was more than the
median, both mentioned parameters were also significantly
correlated, but the relation between population abundance
and the number of captive-reared individuals introduced into
the field in the previous year was negative (N = 19, betawild =
0.434, pwild < 0.0001, SEbetawild = 0.047, betacaptive = 0.286,
pcaptive = 0.0363, SEbetacaptive = 0.124).
6. Discussion
As an important flagship species, the Apollo butterfly has
been the object of successful population recovery projects.
Some projects were aimed at expanding an existing popula-
tion (Dolek and Geyer, 2002; Danková, 1997). One reintro-
duction project led to the restoration of an extinct
population (Lukášek, 2000). An increase in the spatial range
of a metapopulation and increased abundance of wild pop-
ulations have been routinely taken as proof of the effective-
ness of a reintroduction programme (Dolek and Geyer, 2000;
Geyer and Dolek, 1995; Witkowski et al., 1997; Adamski and
Witkowski, 1999a). In the analysed case, close scrutiny of
the effectiveness coefficients (Table 1) yields less optimistic
conclusions. As mentioned earlier, the effectiveness coeffi-
cient calculated as given above is the net population repro-
duction rate determined for a given year and taking into
account captive-reared individuals (Krebs, 1994) In these cir-
cumstances, in order to be deemed stable, the coefficient
for a population should be at least equal to 1. Populations
with a lower rate of reproduction over a period of many
seasons are so-called sink populations, whose survival de-
pends upon an uninterrupted inflow of individuals from
 B I O L O G I C A L C O N S E RVAT I O N 1 4 0 ( 2 0 0 7 ) 1 –7 5

Table 1 – Dispersive activity of butterflies as, percentages of individuals

Year Dispersion inside the metapopualtion centers Dispersion between the centers
E C W G Total

1992 0.0 – – 0.0 0.0 0

1993 2.2 – – 2.0 2.0 0
1994 7.2 – – 5.1 6.3 0
1995 28.6 – – 26.2 27.1 1.2
1996 31.1 32.4 32.1 0.0 31.4 2.4
1997 35.5 34.5 33.7 33.3 34.3 2.1
1998 32.5 32.4 31.6 29.6 32.1 3.9
1999 43.3 40.6 40.5 36.4 41.5 4.7
2000 35.7 37.2 37.0 28.0 36.2 3.8
2001 34.1 32.6 36.6 16.7 34.1 2.9
2002 34.0 31.7 31.5 27.3 32.6 2.1
2003 33.233 35.2 31.4 29.2 33.2 2.5

Table 2 – Effectiveness of the restoration programme, calculated by year

E C W G Total

1992 1.69 – – 1.04 1.67

1993 0.93 – – 0.46 0.84
1994 1.31 – – 0.05 0.13
1995 0.59 – – 0.32 0.48
1996 0.85 – – 0.03 0.36
1997 1.42 1.07 1.06 3.64 1.23
1998 1.76 1.01 0.20 1.80 0.84
1999 1.15 0.28 0.74 0.42 0.69
2000 1.12 0.36 0.44 2.25 0.77
2001 1.06 0.53 0.55 0.23 0.78
2002 0.92 0.68 1.31 1.89 0.96
2003 1.14 1.24 0.51 0.51 0.92

source populations (Pulliam, 1988; Pulliam and Daniellson,
1991), in this case the captive-bredpopulation.
Analyses of the effectiveness coefficients for particular
metapopulation centres give less pessimistic conclusions. In
the Pieniny National Park the Apollo butterfly forms a meta-
population, so not all its parts need have the characteristics
of source subpopulations (Pulliam and Daniellson, 1991).
Moreover, the number of individuals in sink populations
may even markedly exceed the number in the source popula-
tion (Watkinson and Sutherland, 1995).
In the Pieniny metapopulation of the Apollo butterfly, site
E may definitely be recognized as a metapopulation source. It
is inhabited by the strongest and most stable of the Pieniny
Apollo butterfly subpopulations. Since 1996 the net reproduc-
tive rate at site E has either exceeded or been only slightly be-
low 1 (Table 1). In other parts of the metapopulation where
results were calculated, major year-to-year fluctuations were
shown, particularly in the small subpopulation G. This low
stability, possibly the result of low numbers of individuals in
that population, coupled with its peripheral location and its
isolation, means that the role of G in the functioning of the
whole metapopulation is fairly minor.
For the practice of conservation, the relation between pop-
ulation size and the extent of captive-bred supply in previous
years is an important one. Our results show that for a
relatively small population the effect of captive-bred supply
is positive, whereas in abundant subpopulations the opposite
is true. Thus it can be suggested that introducing a large num-
ber of captive-reared individuals to a site may result in local
overcrowding of the population. If, as a result, the combined
numbers of wild and captive-reared populations at a given
site approach the carrying capacity, one can expect a lower
rate of increase due to higher migration (Bujalska and Grüm,
1994) or lower production of fertile offspring (Krebs, 1994). In
the present case it may be assumed that the high number of
imagines translates to a higher number of eggs laid, and
thence a high number of caterpillars emerging at the site in
the next year. Studies completed to date regarded the abun-
dance of stonecrop (Sedum maximum) to be the limiting factor
for the discussed metapopulation (Witkowski et al., 1992,
1997). The emergence of high numbers of caterpillars foraging
on this plant early in the vegetation season may lead to tem-
porary erosion of the food supply. Our oversupply hypothesis
is supported by the finding that the most stable subpopula-
tion, E, is the place where the relative level of captive-reared
supply was lowest. Under such circumstances, supplement-
ing small populations with small groups of captive-reared
individuals should be regarded as the most effective strategy.
Analysis of the movements of individuals between and
within locations indicates that the suggested division of the
Pieniny metapopulation into three distinct centres and one
somewhat isolated site matches its actual dynamics. Within
6 B I O L O G I C A L C O N S E RVAT I O N
the main centres, the coefficient of exchange of individuals
between restored habitat patches was very high; thus these
areas should be treated, at least in part, as population patches
in a greater metapopulation (Harrison, 1991; Harrison and
Taylor, 1997). It might seem surprising that the individual ex-
change coefficients for all the metapopulation centres were
similar, despite the much greater distances between the sites
of the western centre. Straight-line distance, however, is only
one of several factors affecting the probability of migration of
individuals between sites (Hanski, 1997). In the case of the
Apollo butterfly, the presence of open areas between sites en-
abling free dispersion of the species is at least equally impor-
tant (Brommer and Fred, 1999). Studies of Parnassius smintheus
describe the crucial role of habitat structure in the dispersion
activity of the butterflies (Fownes and Roland, 2002; Matter
et al., 2004). This structure involves not only the distribution
of open areas but also the within-meadow structure (Ross
et al., 2005). The lack of suitable migration corridors around
site G is the factor behind the low level of migration among
local individuals, as well as the low number of butterflies
from other sites reaching subpopulation G. Interestingly, in
the studied population the migration ratios of the males
and females did not differ, in contrast to other results and
theoretical predictions (Matter and Roland, 2002). Unfortu-
nately the data available for the Pieniny population are insuf-
ficient to explain this phenomenon.
The exchange of a few percent of the individuals between
the main metapopulation centres seems enough to protect
the metapopulation against genetic erosion (Gilpin, 1996),
but it is not certain whether it is high enough to bring about
spontaneous recovery of a subpopulation if one of the main
centres becomes extinct. At the outset of the recovery pro-
gramme, the Pieniny population of the Apollo butterfly had
an extraordinarily low dispersion coefficient, and raising it
was one of the priorities for the project (Witkowski and
Adamski, 1996; Adamski and Witkowski, 1999a). It seems that
the dispersion coefficient is boosted mainly by introducing
into breeding some individuals from another less isolated
population (Witkowski and Adamski l.c). The drop in long-
distance migration noted in the last three years is most likely
an artefact resulting from the fact that the probability of
recapturing an individual decreases with increased popula-
tion number (Seber, 1982; Turchin, 1998; Adamski, 2004).
In most recovery projects for the Apollo butterfly, the key
activity consists in maintaining or recovering suitable areas.
In the Franconian Alps this activity also involved co-operation
with mining enterprises located in this region (Dolek and
Geyer, 2002). In the Pieniny Mts, introduction of pasture to
habitats suitable for the Apollo butterfly is problematic be-
cause it may conflict with other conservation imperatives of
Pieniny National Park. In such a case, the Apollo butterfly
habitat is maintained through special activities of the Na-
tional Park Service.
7. Conclusions
It can be said that a functional metapopulation of the Apollo
butterfly has been successfully restored in the Polish part of
the Pieniny Mts. Since the supply of captive-reared individu-
1 4 0 ( 2 0 0 7 ) 1 –7
als is continuous, however, it is impossible to determine
unambiguously whether the metapopulation would function
without this kind of support. Undoubtedly the population
patch of the Trzy Korony massif (metapopulation centre E)
can be regarded as a stable part of the metapopulation. Its
net reproduction rate throughout the last six years has ex-
ceeded 1 or fallen only slightly below it (Table 2).
A coefficient for the exchange of individuals between
metapopulation centres amounting to a few percent is too
low to significantly affect the growth rate estimate for any
particular metapopulation centre. The possibility that the sta-
bility of population patch E may be the result of an influx of
migrants from other sites may thus be excluded. Under these
circumstances, this local population patch may be regarded
as a metapopulation source (Pulliam, 1988). Two other centres
seem to have the characteristics of sink populations, but their
actual status can only be determined after the supply of cap-
tive-reared individuals ceases. Also unclear is the status of
the single site at G; significant fluctuations of population
number occur there, but owing to its peripheral position, its
impact on other subpopulations seems very limited. Never-
theless, due to its peripheral position, this population could
function as a stepping stone between the population in the
Pieniny National Park and the more numerous population at
Haligovske Hory in Slovakia.
Acknowledgments
The authors would like to thank Andrzej Kosior and Piotr
Płonka for co-working on the project of Apollo butterfly recov-
ery. Special thanks are due to the staff of the Pieniny National
Park and PIENAP, particularly to Michał Sokołowski, Bogusław
Kozik, Tadeusz Oleś, Štefan Danko and Katarı́na Žlkovanová.
Authors are also grateful Polish and Slovak ministries of envi-
ronment for enabling us exchange of individuals between
captive breadings (Slovak CITES export permission No.
00045/95, Polish CITES import permission No. 4772/96/95).
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