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Entomologica romanica 18: 21-30, 2013 ISSN 1224-2594 / article no.: ER18201304
Differences in adult phenology, demography, mobility and distribution
in two syntopic ecotypes of Maculinea alcon (cruciata vs. pneumonanthe)
(Lepidoptera: Lycaenidae) from Transilvania (Romania)
Natalia , Cristina , Cristian , Alexandra , László
Summary: We present a Mark-Release-Recapture study performed on the populations of two Maculinea alcon ecotypes in a Natura
2000 site from Transylvania, Romania. The Natura 2000 site harbours cultural landscapes with highly biodiverse semi-natural
grasslands, among which several meso-hygrophilous meadows represent the only areas with 4 syntopically occurring European
MaculineaM. alcon. Previous studies have shown that the two M. alcon
distribution. In the case of M. alcon, conservation of the species has to consider the ecological needs of both ecotypes in order to be
meaningful. The unique syntopical occurrence of both ecotypes makes population ecology studies in this area especially important
for providing information for conservation management.
Key words: population ecology, mark-release-recapture, Maculinea alcon ecotypes, syntopic populations.
Introduction
Traditionally managed grasslands in temperate
Europe harbour a high diversity of plants and
of habitats ( et al. 2006). The abandonment or
their fragmentation and isolation has a dramatic
effect on species directly depending on these habitats
( & 2003). The species with most
specialised life cycles have experienced the most
combinations of biotic and abiotic conditions (
et al. 2007).
Maculinea van Eecke, 1915
(synonymised with Phengaris Doherty, 1891 by
et al. 2007) are examples of myrmecophilous species
with a complex life-cycle, adapted to traditional
cultural landscapes that are threatened at European
level ( & 1992, et al. 2005).
After a short period feeding monophagously on their
host plant, fourth instar larvae of Maculinea caterpillars
fall to the ground, where they are adopted by their
host ant species from genus Myrmica Latreille, 1804
( et al. 1998, et al. 1999, et al.
2001). The caterpillars possess a range of adaptations
that enable them to enter and exploit host ant colonies
( & 1992, et al. 1999,
et al. 2009) preying either on ant grubs
(“predatory” Maculinea) or being fed by trophallaxis
(“cuckoo” Maculinea) ( 1968, et
al. 1991, & 1992). Because
Rosidae families) and host ants of the genus Myrmica,
Maculinea
are considered umbrella species and their conservation
et al.
2005, & 2005, et al. 2007,
et al. 2007).
Presently most Maculinea populations are small
and isolated and occupy fragmented habitats, strongly
affected by human activities, mainly because of the
changes in agriculture over the past decades and the
abandonment of traditional land-use (
2001, & 2004, &
2007, et al. 2012).
Maculinea populations have been reported to
be declining in the Carpathian basin and Romania
( 1991, 1993), although, in the latter region, the
precarious economical situation largely maintained
traditional land-use systems with positive effects on
large blue’s habitats ( & 2008, et
al. 2010, et al. 2011). Thus in Romania we can
Maculinea taxa and in
several cases even occurring syntopically: Maculinea
arion Linnaeus 1758, M. teleius Bergsträsser 1779,
M. nausithous kijevensis Sheljuzhko 1928 and M.
alcon with two ecotypes ( et al. 2010,
et al. 2010, et al. 2011, et al. 2012).
where the two forms of M. alcon co-occur: in the
22
Natura 2000 site “Dealurile Clujului Est”, Romania
( et al. 2008, et al. 2011, et
al. 2013).
There are 5 known European Maculinea species,
subspecies or ecotypes. Controversial taxonomical
changes pose problems especially between rebeli and
alcon, and widespread confusion of alcon ecotypes
with taxa xerophila or rebeli
literature (e.g. & 1998).
Because of these taxonomical issues the taxa M.
alcon and M. rebeli are not mentioned on the IUCN
2013 Red list of threatened species.
In this study we aim to analyse ecological
differences and similarities between the two ecotypes
of M. alcon: M. a. ‘cruciata’ and M. a. ‘pneumonanthe’,
occurring syntopically but with distinct host plant
species, Gentiana cruicata and G. pneumonanthe,
in the unique Natura 2000 site from Transylvania
(Romania). But in advance, we need to clarify the
fact that under the name of M. alcon ‘cruciata’ we do
not refer to M. rebeli, M. alcon xerophila or M. rebeli
xerophila. As described by (1904), M.
rebeli is an exclusive alpine species that does not use
Gentiana cruciata as host plant because this gentian
is not found in the type population distribution area
( 2008).
In order to insure the long term survival of these
that shape Maculinea population dynamics and
programs ( et al. 2005a). Thus, we applied
the mark-release-recapture (MRR) method to study
the adult populations of the two Maculinea alcon
ecotypes with the aim of gathering knowledge of
importance for their conservation. The only studies
previously conducted on the two ecotypes in the
et al. 2008) and oviposition behaviour ( et
al.
gather population ecology data about phenology,
demography, mobility and distributions in the two
syntopically occuring ecotypes of M. alcon.
Materials and methods
Study site and species
Our research was carried out in the Natura 2000
site “Dealurile Clujului Est” (Eastern Hills of Cluj),
on a northern exposed meso-hygrophilous meadow of
ca
23.73E, 410-460m a.s.l., Cluj county, Transylvania,
Romania).
The study area is characterized by a mosaic of
two main vegetation types: Molinion caeruleae Koch
1926 in boggy depressions and Cirsio-Brachypodion
Hadac & Klika in Klika & Hadac 1944 in semi-dry
are Molinia caerulea, Sanguisorba ofcinalis,
Gentiana pneumonanthe, Ranunculus polyanthemos,
Carex tomentosa, Juncus conglomeratus, Serratula
tinctoria. For the second alliance, the dominant
and frequent species are: Brachypodium pinnatum,
Bromus erectus, Asperula cynanchica, Carex humilis,
Polygala major, Cirsium pannonicum, Centaurea
scabiosa, Ranunculus polyanthemos, Prunella
grandiora, Gentiana cruciata, Veronica teucrium,
Trifolium pannonicum, T. ochroleucon and others
( et al. 2011). The pasture was abandoned
progressively after 1990 and in 2011 was partially
mown (3% of the entire study area) ( personal
communication). In the period 2009-2012 the study
area was partially and intensively grazed by sheep.
Maculinea species
cohabit syntopically: M. alcon (with two ecotypes),
M. teleius and M. nausithous kijevensis ( et al.
2011, et al. 2013). The females of one of the
M. alcon ecotypes oviposit on Gentiana cruciata,
the other on Gentiana pneumonanthe. Since the
taxonomy of MaculineaG.
cruciata is not clear ( et al. 2006,
2008), we prefer to call the two ecotypes according to
their host plants, thus: the form using G. cruciata is
referred to as M. a. ‘cruciata’, while the form utilising
G. pneumonanthe as M. a. ‘pneumonanthe’.
The two gentians used by M. alcon ecotypes as host
plants, co-occur in some patches, but G. pneumonanthe
has higher abundance in wet depression with Molinia
caerulea, while G. cruciata is more abundant in semi-
dry areas.
For M. a. ‘pneumonanthe’ there is strong evidence
that the host ant species is Myrmica scabrinodis
( et al. 2008, et al. 2013). The M.
a. ‘cruciata’ caterpillars were found (G. cruciata
patches) in Myrmica schencki and M. sabuleti nests
( et al. 2008).
Sampling method
The two syntopic populations of M. alcon
mark-release-recapture (MRR) method. The MRR
method was applied in 2010 on M. a. ‘pneumonanthe’
population and in 2011 both on an M. a. ‘cruciata’
and an M. a. ‘pneumonanthe’ populations. Qualitative
were registered in the years 2009-2011.
every second or third day (with exceptions caused by
unfavourable weather) between 9:00 A.M. and 7:00
years between the 11 June – 9 August in 2010 and
2011. In the case of each capture/recapture, gender and
plant, sitting on food plants, oviposition, mating) of
the captured individual, capture time and coordinates
23
of the exact location were recorded. Coordinates were
recorded with a GPS device Garmin GPSmap 60 CSx.
In order to map the host plants of the two ecotypes:
Gentiana cruciata and Gentiana pneumonanthe, a grid
of squares of 50x50 m was established that covered
the grid was investigated and all individuals of the
two plants were marked with a GPS device (Garmin
GPSmap 60 CSx) in June and July both years 2011
and 2012.
Statistical analyses
MRR data from the years 2010 and 2011 was
analyzed to estimate the current population size,
M.
a. ‘pneumonanthe’ and M. a. ‘cruciata’ ecotypes.
Data was analysed separately for each ecotype with
the Cormack-Jolly-Seber type constrained models
( & 1996, &
1999) using the program MARK 6.0 package (
& 2010). The performances of the models
were assessed with the Akaike Information Criterion
corrected for small sample size (AICc) (
1973, & 1989). As recommended
by & (2001), after running
number of parameters.
Individual life span was estimated using the
formula ê=(1–φ)–1–0.5 (ê is the individual life span,
φ is the survival probability) ( et al. 2005b).
The parameters resulted from analysing the data
with the program Mark 6.0, survival and capture
probability, were used to estimate daily number of
individuals in each capture occasion (i.e. days of
capture) and recruitment of new individuals into the
population and the total number of individuals in the
two populations (of the two ecotypes).
We compared the number of marked individuals,
both years between the two ecotypes with a Mann-
Whitney U-test (with the program Past 2.09,
et al. 2001), because data did not follow a normal
distribution.
Flight distances were calculated separately for
males and females within season as the distances
distances from same day recaptures for both sexes and
ecotypes using program ArcMAP 9.2. Comparisons
sexes were also computed with a Mann-Whitney
U-test (with the program StatView 5.0), because
datasets were not normally distributed. A Chi squared
www.quantpsy.org/chisq/chisq.htm) on the seasonal
evenly distributed on three and four, respectively,
distance classes (0-100 m, 101-200 m, 200-300 m,
over 300 m). For M. a. ‘cruciata’
we could not perform this test because sample size
was too small.
In order to compare the spatial distributions of the
G. cruciata and G. pneumonanthe plants and of the
adults of M. a. ‘cruciata’ and M. a. ‘pneumonanthe’
we overlaid a grid of 20x20 m over the map of
Quantum GIS 2.2.0 - Valmiera. Of these we randomly
selected 4 times 25 squares and counted the number
Correlation between the distribution of host plants
using Spearman Rank Correlation with the program
Past 2.09 ( et al. 2001).
Results
Phenology and population parameters
M. a. ‘cruciata’
between the 14th and 26th of June in 2011. Previous
observations from the years 2009 and 2010 showed
Figure 1. Marked individuals of M. a. ‘cruciata’(grey bars – data from 2011) and M. a. ‘pneumonanthe’ (empty bars – data
from 2010, dark bars – data from 2011) ecotypes in the investigation area Fanatul Domnesc (Cluj County, Romania).
24
30th June at the latest. M. a. ‘pneumonanthe’
approximately 4 weeks, between the 11th of July and
the 7th of August in 2010 and 2011 (Fig. 1).
In 2010 330 individuals (204, 62% males and 126,
38% females) of M. a. ‘pneumonanthe’ were captured
and marked. Of these 40 (12.1%) individuals (29
males and 11 females) were recaptured at least once
on a different capture occasion.
In 2011 294 individuals (225, 77% males and
69, 23% females) of M. a. ‘pneumonanthe’, and 165
individuals (112, 68% males and 53, 32% females)
of M. a. ‘cruciata’ were captured and marked. Of
these 20 (6.8%) individuals (16 males and 4 females)
of M. a. ‘pneumonanthe’, and 16 (9,7%) individuals
(13 males and 3 females) of M. a. ‘cruciata’ were
recaptured at least once on a different capture occasion
(Table 1). The number of marked individuals did not
Whitney U-test, p>0.05).
For M. a. ‘cruciata’ a population of 1073
individuals was estimated in the year 2011 with the
model Phi(.) p(t) (i.e. the model showing constant
survival probability and different capture probability
over time) (Table 2). For M. a. ‘pneumonanthe’ a
population of 1277 individuals was estimated for the
year 2010 with the model Phi(.) p(g) (i.e. the model
showing constant survival probability and different
capture probability for males and females) and a
population of 1296 individuals for the year 2011 with
the model Phi(t) p(g) (i.e. the model with different
survival probability in time and different capture
probability for males and females).
The survival probability (Phi) was 0.85 (±0.09
SE) for M. a. ‘cruciata’, and 0.71 (±0.04 SE) and an
average of 0.54 (±0.34) for M. a. ‘pneumonanthe’ in
2010 and 2011 respectively. The capture probability
(p) was on average 0.21 (±0.28 SE) for M. a. ‘cruciata’,
and 0.26 (±0.11 SE) and 0.33 (±0.22 SE) for M. a.
‘pneumonanthe’ in 2010 and 2011 respectively.
Individual life span was calculated at 6 days for M.
a. ‘cruciata’ in 2011 and at 3 and 2 days respectively
for M. a. ‘pneumonanthe’ in 2010 and 2011 (Table 2).
Daily population estimates (Ni) for each capture
years for M. a. ‘pneumonanthe’ and between ecotypes
in 2011 (Mann-Whitney U-test: p>0.05) (Fig. 2).
With the exception of M. a. ‘pneumonanthe’ males in
2010 (unimodal pattern), for both ecotypes the adult
Bi) have well-
is similar between years for M. a. ‘pneumonanthe’
and also between ecotypes in 2011 (Mann-Whitney
U-test: p>0.05).
Figure 2. Within-season daily population size estimates (Ni). Empty bars –M. a. ‘pneumonanthe’ 2010, dark bars - M. a.
‘pneumonanthe’ 2011 and grey bars – M. a. ‘cruciata’ 2011.
Year Ecotype Marked individuals Recaptured Individuals* Recapture
Ratio (%)
Males Females Total Males Females Total
2010 M.a.’p’ 204 126 330 29 11 40 12.12
2011 M.a.’p’ 225 69 294 16 4 20 6.80
2011 M.a.’c’ 112 53 165 13 3 16 9.69
*individuals recaptured at least once on a different capture occasion in the same study site
Table 1. Summary of the MRR study on Maculinea alcon ‘pneumonanthe’ (M.a.’p’ ) and M. alcon ‘cruciata’ (M.a.’c’ ) in
25
Generally, captured individuals and also estimates
show a clear male–biased ratio. The sex ratio differed
2011 for both ecotypes (Table 2).
Mobility and distribution
between consecutive recapture dates) for M. alcon
‘cruciata’ was 100 m, and 97 m and 75 m for M. a.
‘pneumonanthe’ in 2010 and 2011 respectively. The
distances from same day recaptures were similar:
121 m for M. alcon ‘cruciata’ in 2011 and 97 m and
110 m for M. a. ‘pneumonanthe’ in 2010 and 2011
and between ecotypes (Mann-Whitney U-test p>0.05
registered (distance between consecutive recapture
dates) was 345 m for a female M. a. ‘cruciata’.
than 200 m in both ecotypes and the Chi squared
2=34.9,
DF=2, p=3x10-8
200 m for M. a. ‘cruciata’ and up to 100 m for M.
a. ‘pneumonanthe’
followed the same unevenly distributed pattern on
distance categories with most distances under 200 m
for M. a. ‘pneumonanthe’ 2=63.1, DF=3, p=0).
When comparing the spatial distribution of
the adults and host plants for both ecotypes we
M.
a. ‘pneumonanthe’ and G. pneumonanthe (rs=0.67,
p=1.7x10-10) and a lower correlation between M. a.
‘cruciata’ and G. cruciata (rs=0.30, p=0.003). No
other correlations were found between distributions
of the two plants, between the distributions of the
two ecotypes and the distributions of plants and
irrespective ecotypes.
Figure 3. Within – season recruitment (Bi) in the populations
investigated. Broken line represent total recruitment
between consecutive capture days, squares – males and
circles – females.
Year Ecotype Model
Seasonal
population
size
Sex ratio
(males:females)
%
φ
pLifespan
(days)
Males Females
2010 M.a.’p’ Phi(.) p(g) 1313 52:48 0.712 0.32 0.20 2.98
2011 M.a.’p’ Phi(t) p(g) 1297 73:28 0.461 0.36 0.29 2.05
2011 M.a.’c’ Phi(.) p(t) 1073 68:32 0.845 0.21 0.21 5.98
Table 2. Basic parameters of the Cormack-Jolly-Seber model (p average daily capture probability; φ average daily survival
rate), population estimates (computed with MARK 6.0 program), sex ratio and individual life span in Maculinea alcon
‘pneumonanthe’ (M.a.’p’ ) and M. alcon ‘cruciata’ (M.a.’c’ ) populations,
County, Romania).
26
Discussion
Phenology and population parameters
Our phenology results show that the two ecotypes
periods.
Considering our phenological data, the two
ecotypes from Transylvania were clearly separated
in each year by at least two weeks. Phenological
phenology of the two host plants.
The possibility of the two ecotypes being two
generations of the same species has to be excluded in
this case, as M. a. ‘cruciata’ ecotype needs on average
11-23 months ( et al. 1998, et
al. 2000, & 2007,
unpubl. data) and exceptionally (in optimal laboratory
conditions) at least 6 weeks ( 2005) for the
preadult development. Furthermore, &
(2007) found that only a 3 month period
of cooling triggered the completion of development
(i.e. an obligatory diapause) in both ecotypes.
Considering the population parameters, the two
ecotypes were similar in population size, survival,
capture probability and recruitment. Only individual
life span was considerably higher in M. a. ‘cruciata’,
with 6 days, compared to M. a. ‘pneumonanthe’ with
only 3 days.
2011 was characterised by lower recapture rates.
estimates in both ecotypes. However when population
parameters of 2011 were compared to those of 2010
for M. a. pneumonanthe
were found. We have thus used these estimates in
order to check for possible differences between
population parameters of ecotypes, as the work and
analysis method was the same for both.
Both ecotypes showed a skewed sex ratio and a
bimodal recruitment in the year 2011.
et al. (2005) suggested that a deviation
from the expected sex ratio can be traced down to the
higher recapture rates of males, due to the fact that
mobile females. Females, on the other hand, tend to
the undergrowth and may be easily overlooked.
After et al. (2005a, 2009) the bimodal
period (Bi) indicates the presence of biannual
larvae. Biannual larvae were found in all European
Maculinea species: M. rebeli ( et al. 1998),
M. alcon, M. arion ( et al. 2000), M.
teleius and M. nausithous ( et al. 2006). Even
though our data shows a bimodal recruitment pattern
within season for both ecotypes, we found no larval
Myrmica
colonies ( unpublished data). Also, larval
development in laboratory experiments with ecotypes
from the same area, never indicated polymorphic
growth rates ( unpublished data), in contrast to
developing larvae ( et al. 2000,
et al. 2006).
Mobility and distribution
An increasing number of studies showed that
host plant development rather than host-ant presence
et al. 1997, et al. 2005c, &
the distribution of the respective host plant. M. a.
‘pneumonanthe’ showed a higher density in patches
of G. pneumonanthe resulting in a higher correlation
M. a. ‘cruciata’ was more dispersed, similarly to
the host plant distribution, G. cruciata, resulting in a
two distributions. At the same time, distributions of
the two host plants did not overlap and correlations
between ecotype and irrespective host plant were not
found.
& (2004)
presented the hypothesis that M. alcon ([Denis &
Schiffermüller]1775) could also use G. cruciata apart
from G. pneumonanthe as an additional host plant,
by studying oviposition preferences. But their study
could not prove that eggs laid on G. cruciata could
sustain viable offspring, because only one larva was
found in ant nests in the vicinity of G. cruciata plants.
Also the study of et al. (2013), conducted
Romania), showed clearly separated oviposition
behaviour and preference in the two ecotypes for their
respective host plant.
sedentary. Their limited mobility and distribution
overlapped with that of their host plant indicates that
two ecotypes of M. alcon
Domnesc (Cluj County, Romania).
27
they establish home - ranges, not moving far from
their emergence site. This phenomenon has been
reported also in other recent MRR studies for other
Maculinea species ( & 2008,
et al. 2008, &
2009, et al. 2011, et al. 2013).
& (2008) suggested that
keeping close to the place of eclosion is an adaptation
of Maculinea
The establishing of home-ranges of Maculinea
A low number of emigrants increases the risk of
extinction of declining populations, and affects
the colonisation process dynamics ( et al.
2005a). Even though on rare occasions Maculinea
in colonizing new areas, the probability of such an
event is probably lower than the local extinction rate
( et al. 2005a).
Implications for conservation
Meadows that shelter more than 2 Maculinea
species are very rare. The habitats in the Natura
2000 site ‘Dealurile Clujului Est’ are probably
unique in Europe, because in some northern exposed
meso-hygrophilous meadows of this site four of the
European Maculinea taxons co-habit syntopically:
Maculinea arion, M. alcon ‘pneumonanthe’, M.
alcon ‘cruciata’ (also on southern exposed sites),
M. teleius and M. nausithous kijevensis. Such places
are particularly fragile because of each Maculinea
their conservation a hard task.
The presence and preservation of vigorous
Maculinea populations in the area has been fostered
over time through a randomized mowing regime of
the parcels owned by different farmers. In this system,
every year, a part of the parcels which harbour
fragments of Maculinea populations remain unmown
or are mown very late in summer, maintaining in this
way the metapopulational structure of the species and
having as a result the preservation of the large blues
( et al. 2011).
Their reduced mobility, home-range behaviour and
distributions closely connected to that of their host
plants all lead to the greater susceptibility to isolation
of populations through reducing their probability to
disperse and colonize new habitats in the two analysed
M. alcon ecotypes. Thus, habitat fragmentation by
land use increases the vulnerability of M. a.’cruciata’
and M. a. ‘pneumonanthe’.
The cultural landscapes and mosaics of secondary
habitats resulted from differently used meadows in the
is already threatened by factors such as short- and
long-term abandonment of traditional land-use (hand-
grazing (especially with sheep), drainage works and
local urban development plans ( et al. 2010,
et al. 2011, et al. 2011).
most threatening activities are abandonment and
intensive grazing. The abandonment of mowing and
grazing is likely to have a positive short-term impact
on Maculinea populations ( et al. 2011). The
species diversity in general is supported by the
studies of & (2000), et
al. (2005), & (2007), and
& (2011). However, after 4-5 years, this
process will most likely go into reverse and cause the
decline of biodiversity & 2007),
affecting the Maculinea populations as well. The
same, abandonment of heterogeneous mowing of
small meadow plots potentially leads to an alteration
of the metapopulational structure of Maculinea spp
( N. unpubl. data).
populations in abandoned hay meadows depend on
the intensity and type of grazing. In general, extensive
grazing with cattle can have a positive impact because
it prevents or slows down the secondary succession
towards shrubs and can therefore contribute to
For M. a. ’cruciata’ and M. a.’pneumonanthe’ in
particular, even an extensive grazing has a negative
impact: host plants with eggs can be easily eaten or
trampled by sheep ( A. unpublished data).
In order to protect the species diversity and the
viable metapopulational structures, there is a strong
need to develop strategies for maintaining the current
mosaic of habitats characterized by different stages
of succession ( et al. 2005, et al.
2007, et al. 2011), e.g. through more research,
support of traditional small-scale farming and active
nature conservation management.
Many studies emphasize the importance of
maintaining an interconnected network of suitable
habitats ( 1999) but also of habitat quality,
especially for sedentary species ( et al.
2001, & Van Dyck 2005, et al. 2007).
Conservation of the two M. alcon ecotypes should
aim at improving habitat quality. Considering their
low mobility, the spread of the local populations can
distance (i.e. 200 m). Habitat quality can be improved
like extensive mowing once a year, either few weeks
Within an interdisciplinary project funded by the
Deutsche Bundesstiftung Umwelt (DBU; project
number 27559) a pilot agri-environment programme
was carried out in 2011 and 2012. This project offered
incentives to apply the earliest mowing on the 25th of
August and tractor and hand mowing was allowed.
The short-term outcomes of this project showed a
28
with the lower quality of hay, but on the other hand,
most farmers were encouraged by courses and regular
support, so that they willingly cooperated in the
project ( et al. 2012).
Maculinea
the reduction in mowing earlier in the season. Thus, in
2012 a new package of the national agri-environment
scheme was introduced for a restricted geographical
genus Maculinea: Agri-environment package 6
Maculinea
spp”. This package aims at maintaining high habitat
quality for Maculinea
Euros/ha) can be received for all permanent grasslands
and the main requirements are: the earliest mowing
date on the 25th of August, mowing only allowed by
scythe or small hand mowing machines, grazing with
min 0.3 livestock unit/ha (0.3 cows /ha or 1.8 sheep/
ha) and max. 0.7 livestock unit/ha (0.7 cow per ha or
4.2 sheep/ha) (Axa II, www.apia.org).
This might be the beginning of long-term education
for agri-environment conservation in Romania;
however the long-term effects of this measure cannot
be estimated yet. The effective implementation of
such agri-environment measures often depends on
management of local administration and willingness
of stakeholders to apply for incentives and practice
the recommendations of the measures after NGO
support and educational programs are reduced.
Acknowledgements
distribution, to I. Paulini for her help with important
management information from the studied area and
to Z. Varga and M. Davies for important comments
on the manuscript. This work was granted from the
Sectoral Operational Programme for Human Resources
Social Fund, under the project number POSDRU
89/1.5/S/60189 with the title “Postdoctoral Programs for
Sustainable Development in a Knowledge Based Society“
to C. Craioveanu.
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Natalia
(contact author)
Department of Taxonomy
and Ecology,
Clinicilor 5-7,
400006 Cluj-Napoca,
Cluj County, Romania
nataliatimuss@gmail.com
Received: 10.12.2013
Accepted: 22.12.2013
Published online: 28.12.2013
Published: 30.01.2014
Online article number: ER17201305
Alexandra
Department of Taxonomy
and Ecology,
Clinicilor 5-7,
400006 Cluj-Napoca,
Cluj County, Romania
Cristina
(contact author)
Department of Taxonomy and
Ecology,
Clinicilor 5-7,
400006 Cluj-Napoca,
Cluj County, Romania
cristinacraioveanu@gmail.com
Cristian
Department of Taxonomy
and Ecology,
Clinicilor 5-7,
400006 Cluj-Napoca,
Cluj County, Romania
László
Department of Taxonomy
and Ecology,
Clinicilor 5-7,
400006 Cluj-Napoca,
Cluj County, Romania
