Timuş N. PhD summary 2014. Taxonomy, biology and ecology of myrmecophilous butterfly Maculinea teleius (Lepidoptera, Lycaenidae) from Cluj and Dej Hills Area (Cluj County)

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Cluj-Napoca
2014
Taxonomy, biology and ecology of
myrmecophilous butterfly Maculinea
teleius (Lepidoptera, Lycaenidae) from
Cluj and Dej Hills Area (Cluj County)
“Babeş-Bolyai” University
Facultaty of Biology and Geology
Department of Taxonomy & Ecology
PhD student
Natalia Timuş
Scientific Advisor
Prof. Dr. László Rákosy

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“Babeş-Bolyai” University
Facultaty of Biology and Geology
Department of Taxonomy & Ecology
PhD student:
Natalia Timuş
Scientific Advisor:
Prof. Dr. László Rákosy
Summary of the PhD Thesis
Taxonomy, biology and ecology
of myrmecophilous butterfly Maculinea
teleius (Lepidoptera, Lycaenidae) from
Cluj and Dej Hills Area (Cluj County)
Cluj-Napoca, 2014

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Content
Introduction 3
Study area 4
Chapter 1. General aspects of ecology and taxonomy in Maculinea teleius 5
Chapter 2. The adoption process of Maculinea caterpillars by Myrmica scabrinodis ants
infested with ectoparasitic fungus Rickia wasmannii versus non-infested 11
Chapter 3. Survival and development of Maculinea caterpillars in Myrmica scabrinodis
colonies infested with Rickia wasmannii versus non-infested 14
Chapter 4. Demographic parameters of Maculinea teleius and M. nausithous kijevensis
from “Dealurile Clujului Est” 18
Chapter 5. Mobility and spatial distribution within habitat of Maculinea teleius and
Maculinea nausithous kijevensis butterflies 21
Chapter 6. The land-use impacts on M. teleius populations from Natura 2000 site
“Dealurile Clujului Est” 25
Selected references 28
List of publications 32
Acknowledgements 36
Key words: Maculinea, Myrmica, Laboulbeniales, Ichneumonidae, myrmecophily,
social parasites, parasitoids, morphology, ecology, taxonomy, biological
communities, mark – release – recapture (MRR), population ecology, local
specialisation, polymorphic larval growth, metapopulation system, source – sink
dynamics, traditional land-use, conservation, Transylvania, România.

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Introduction
The butterflies of genus Maculinea Van Eecke 1915 (Lepidoptera,
Lycaenidae) are extremely specialised myrmecophilous species with a complex life-
cycle, that involves diferent species of plants ásteaeae şi ‘osaeae and species
of ant from Myrmica Latreille 1804 genus (Elmes & Thomas 1992, Fiedler 1998).
The Large Blues are examples of species adapted to traditional cultural
landscapes that are threatened at European level (Elmes & Thomas 1992, Grill et
al. 2008, van Swaay et al. 2012). In Romania the main factors for the decline or the
extinction of some populations of Maculinea species are the habitat
fragmentation, long-term abandonment of traditional land-use (hand-mowing,
extensive grazing etc.), the inappropriate period of mowing (in the flight season of
Maculinea), intensification of grazing (especially with sheep), drainage works, the
conversion of grasslands to arable land and local urban development plans (Vodă
et al. , Tiuș et al. , ‘kos , Tiuș et al. a).
Maculinea butterflies are considered umbrella species and their
conservation benefits many other threatened species (Thomas et al. 2005, Skórka
et al. 2007, Anton et al. 2007). Also they ae egaded as the flagships of
biodiversity conservation in Europe (Thomas and Settele 2004, Kühn et al. 2005),
epitomising the way that the management of their protection goes hand in hand
with the preservation and conservation of human culture (Grill et al. 2008).
Up until 2009 in Romania, apart from limited faunistical data, little was
known about any of the Maculinea taxa. The Large Blues butterflies have started
being thoroughly studied after sites with Maculinea taxa from Dealurile Clujului
Est area were dicovered by Prof. Dr. László Rákosy. The habitats in the Natura
 site Dealuile Clujului Est ae poal uiue in Europe, because in some
northern exposed meso-higrophilous meadows five of the European Maculinea
taxa cohabit syntopically: Maculinea arion Linnaeus 1758, M. alcon
pneumonanthe Denis & Schiffermüller 1775, M. alcon cruciata (also on
southern exposed sites), M. teleius Bergsträsser 1779 and M. nausithous kijevensis
Sheljuzhko 1928 (Rákosy 2013, Tiuș et al. 2013a). This was the reason why in
2009 a series of biological and ecological studies, such as mark-release-recapture
studies, Myrmica nests investigation, laboratory experiments etc., were initiated.

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Study area
Our studies were carried out in Natura 2000 site Dealurile Clujlui Est. Study sites:
1. meso-higrophilous abandoned meadow of cca 41 ha named Fațul
Domnesc (Chapters 1, 2, 3, 4, 5 and 6)
2. meso-higrophilous hay meadow of cca 92 ha named Fațul “ătes
(Chapters 4, 6)
3. meso-higrophilous hay meadow of cca 81 ha named “eheliște (Chapter 6)
Fig.1. Map of „Dealurile Cluj Est”, Natura 2000 site, with the main categories of land-use in
2012 (modified after the map created in the project „Elaborarea planului de management
integrat pentru situl de importanță comunitară ROSCI0295 - Dealurile Clujului Est”, Romanian
Lepidopterological Society)

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Chapter 1. General aspects of ecology and taxonomy in
Maculinea teleius
Maculinea teleius life cycle
In Romania the Scarce Large Blue (Maculinea teleius) butterfly is on the
wing from July till the end of August. Females oviposit [Fig.1.1 (1)] on the young
inflorescences of Sanguisorba officinalis, the only host plant used by these
butterflies (Malicky 1968, Thomas 1984). During the first three instars (3-4 weeks)
caterpillars will stay on the flowerheads and feed monophagously on the
developing seeds of the host plant (Thomas 1984).
Immediatly after the last moulting [L3 exuvia Fig.1.1(2)], fourth instar
caterpillars of M. teleius descend on the ground with the help of a silk filament
(Fig.1.1 (3)] secreted from specialised glands. The first interaction with host ants
evolves into an adoption ritual that varies depending on Maculinea and Myrmica
species. In the case of M. teleius adoption process can last from tens of minutes to
several hours, in which time the caterpillars are palpated by the ants with their
antennas. In response to antennation, caterpillars offers droplets secreted from
dorsal nectary organ (DNO) (Malicky 1968). The ants eagerly imbibe each droplet
[Fig.1.1(4)], then will carry the caterpillars to the their nest. [Fig.1.1(5)] (Frohawk
1924, Thomas 1984).
While living in Myrmica nests, Maculinea caterpillars feed in two different
ways: caterpillars of M. reeli =cruciata ecotype) and M. alcon are fed directly
by nurse ants via trophallaxis ad ae teed ukoo speies Eles et al. ,
whereas caterpillars of M. teleius, M. nausithous and M. arion prey on ants brood
ad ae alled pedato speies Thoas & Wardlaw 1992).
The caterpillars hibernate in host ant nests (Thomas & Wardlaw 1992). In
spring (after 10/23 months, Witek et al. 2006) M. teleius caterpillars grow rapidly
[Fig.1.1(6)] (Witek et al. 2011) till pupation [Fig.1.1(7)], wich takes place in the
uppe haes of the ats ests alled solarium [Fig.1.1(8)]. The adults eclose
after 2 weeks [Fig.1.1(9)] (Thomas 1984, Witek et al. 2006).
Adaptations to myrmecophily of Maculinea species
The fourth instar caterpillars possess a range of morphological,
physiological, chemical and behavioural adaptations that enable them to enter and
exploit Myrmica host ant colonies. Some of them are described below alongside
with our own observations. Maculinea caterpillars have a thick and tough cuticle.
As has been demonstrated by Malicky (1970), the cuticle of lycaenid caterpillars is
5-20 times thicker than that of other lepidopteran caterpillars of comparable size.

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Fig.1.1. Maculinea teleius life cycle.
The Maculinea caterpillars can also retract their head under their
prothoracic shield (Fig.1.2.a) – a cuticular fold of T1 (thoracic segment 1) (Fiedler
1991, ”liińska et al. . Thus, the most vulnerable organs (nervous system) are
well protected against possible ant-attacks (Fiedler 1991) (Fig.1.2.b). When
feeding on ant larvae, the M. teleius caterpillars pull the prothoracic shield over
the head, thereby hiding their pray.
The predatory caterpillars of M. teleius (LIV) have on dorsal surface long
and thick setae disposed regularly in pairs with 3/2 setae on thoracic segments and
a single pair of setae on abdominale segments (AII – AVII) (Fig.1.2a). M. nausithous
kijevensis caterpillars always have on the dorsal surface 4 long and thick setae
disposed in pairs on thoracic segments T2 and T3 (pers. obs.) (Fig1.2b). The
presence of the two pairs of long setae could be a specific morphological trait of
M. nausithous kijevensis caterpillars, differentiating it from M. nausithous species.
Studies are needed for confirmation.
During the adoption process, M. teleius caterpillar contracts the first
abdominal segment (AI) (Fiedler 1990), which is smaller and without dorsal setae
(Fig.1.2.b). This feature enables the ant to use its mandibles in order to pick up the
M. teleius caterpillar and carry it into the nest.

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The Maculinea caterpillar emanates sounds from muscular contractions in
the abdomen or from sclerotized structures (Fig.1.2.d) between segments 4 and 7
(Barbero et al. 2009). Sounds produced by pupae and larvae of the butterfly
Maculinea mimic those of queen ants more closely than those of workers, enabling
them to achieve high status within ant societies (Barbero et al. 2009).
Over the dorsal surface of Maculinea caterpillar are scattered small
epidermal glands called Pore Cupola Organs (PCOs) (Fi.1.2.f). PCOs secrete
substances that generally may serve to pacify aggressive ants (Malicky 1970,
Fiedler 1991).
The dorsal nectary organ (DNO) is located on the seventh abdominal
segment (Fig.1.2.e,f) and is a specialised exocrine gland that produces nutritious
secretions (Fiedler 1991). The DNO secretes droplets when stimulated by ants via
antennation (Malicky 1970).
The DNO of M. teleius and M. n. kijevensis caterpillars (LIV) is surrounded
by a field of dendritic setae (pers obs.) (Fig. 1.2.f). It is highly likely that these are
specialized setae with mecanoreceptive properties as was described for
Polyomatus icarus caterpillars which dendritic setae are able to preserve the exact
time pattern of tactile stimulation of a specific species of ants (Tautz & Fiedler
1992). All our attempts to elicit the DNO secretion of M. teleius caterpillars (tactile
stimulation with human hairs, brushes, amputated ant heads) were unsuccessful.
Only live ants antennation during adoption procces could obtain the DNO
secretion. Probably the mechanosensory hairs of M. teleius and M. n. kijevensis
caterpillars can recognise the antennation of Myrmica host ants.
The PCOs and mechanosensory hairs are more dense in M. nausithous
kijevensis caterpillars then in M. teleius.
Taxonomy aspects
The genus Maculinea Van Eecke 1915, Lycaenidae family, Polyommatinae
subfamily, comprises species restricted to the Palaearctic region (Sibatani et al.
1994, Wynhoff 1998).
Recent publications based on both molecular and morphological data (Als
et al. 2004, Pech et al. 2004, Fric et al. 2007), have shown that species of
Maculinea and Oriental genus Phengaris form a monophyletic group and according
to Fric et al. (2007) Maculinea Van Eecke, 1915 should be considered a junior
subjective synonym of Phengaris Doherty, 1891. Due to the widespread usage of
the name Maculinea (references in Barbero et al. 2012) some of the authors have
asked the International Commission on Zoological Nomenclature to conserve the
name Maculinea against Phengaris. It was due to the folowing reasons that I used
the name Maculinea in the doctoral thesis: the decision by the ICZN is still pending,

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in Romania a new package of the national agri-environment scheme was
introduced in 2012: Agri-eioet Pakage  Gasslads ipotat fo
butterflies esp. Maculinea spp.
Cryptic speciation has been hypothesised for Maculina teleius based on
divergent mtDNA sequences (Als et al. 2004, Fric et al. 2007, Ugelvig et al. 2011).
According to recent sudies (Ritter et al. 2013) based on mtDNA barcoding, nuclear
microsatellite analyses and Wolbachia screening, hypothesis of cryptic speciation
within Maculinea (Phengaris) teleius is rejected. The major splits in the mtDNA
phylogeny can be explained by Wolbachia infections. Furthermore, the geographic
isolation during Pleistocene glaciations, which likely took place in Central or
Eastern Asia, contributed to differentiation of mitochondrial and nuclear genomes
(Ritter et al. 2013). The phylogeographic hypothesis proposed by Ritter et al.
(2013) is also corroborated by the fact that all described subspecies in M. teleius
are restricted to the Eastern Palaearctic (M. t. sinalcon Murayama 1992, M. t.
obscurata Staudinger 1892, M. t. euphemia Staudinger 1887) and mainly to Japan
(M. t. hosonoi Takahasi 1973, M. t. kazamoto Druce 1875, M. t. ogumae
Matsumura 1910 and M. t. daisensis Matsumura 1926) (Sibatani et al. 1994).
Other taxa recorded in Maculinea-Myrmica system
In the study area (Dealurile Clujului Est) the Myrmica scabrinodis Nylander
1846 ant species is the main host for the Maculinea taxa, and for the syrphid
Microdon myrmicae Schönrogge et al. 2002 (Fig. 1.3.a,b,c). Also these ants are
parasitised by ectoparasitic fungus Rickia wasmannii Cavara 1899 (Tartally et al.
2007) (Fig.1.3.d,e).
In Romania there are limited available data relating to Maculinea
parasitoids species. Pupae belonging to the two Maculinea alcon ecotypes
parasitised by Ichneumon eumerus ee foud at Șadu ad ‘ăsui (Tartally
2008). As a result of investigations carried out on Myrmica nests in 2010 and 2012
Luna de Jos, Cluj county, Transylvania, Romania), M. alcon „peuoathe
ecotype) pupae were found in three Myrmica scabrinodis colonies. In total 13
infested pupae (Fig.1.3.g) were collected and kept under laboratory conditions,
from which 3 males and 5 females Ichneumon balteatus Wesmael 1845 (Fig.1.3.h)
emerged after 1-2 weeks. Prior to our recent discovery, Ichneumon balteatus was
only known to utilise two host species: Melitaea cinxia (L., 1758) (Nymphalidae)
and Calliteara pudibunda L. (Lymantriidae) (Constantineanu 1959, Paul & Hanski
2004), and Ichneumon eumerus was the only recorded parasitoid of M. alcon
butterflies (Shaw et al. 2009). Therefore the association Maculinea alcon and
Ichneumon balteatus is new to science Tiuș et al. .

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Fig. 1.2. SEM photo Ciprian Mihali
a) Dorsal setae in Maculinea teleius caterpillar (LIV), PS - prothoracic shield. b) Dorsal setae in M.
nausithous kijevensis caterpillar (LIV). c) The first abdominal segment (AI) of Maculinea teleius
caterpillar. d) Detail - sclerotized structures between abdominal segments, possible stridulatory
organ in M. nausithous kijevensis caterpillars. e) Dorsal view of Maculinea teleius caterpillar, DNO
– dorsal nectary organ. f) Mechanosensory hairs (yellow), Pore Cupola Organs (red), DNO –
dorsal nectary organ.

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Fig. 1.3. Photo: Natalia Timuş, SEM photo: Lucian Barbu Tudoran
a) Microdon myrmicae larva (LIII) and Myrmica scabrinodis ants. b) Microdon myrmicae
pupa. c) The eclosion of Microdon myrmicae adult, 2012 May, Fânaţul Domnesc, (Luna de
Jos, jud. Cluj). d) Myrmica scabrinodis queen infested with Rickia wasmannii. e) Rickia
wasmannii thallus on Myrmica scabrinodis. f) Ventral view of Maculinea alcon
“pneumonanthe” pupa, arrows indicate the butterfly antennae. g) M. alcon “pneumonanthe”
pupa (lateral view) infested with Ichneumon balteatus, the arrow indicate the parasitoid
antennae. h) Ichneumon balteatus male.

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Chaper 2. The adoption process of Maculinea caterpillars by
Myrmica scabrinodis ants infested with ectoparasitic fungus
Rickia wasmannii versus non-infested
Timuș N., Csata E., Witek M., Babik H, Czekes Z., Erős K., Rákosy L., Markó B.: Parasitic
fungi as key to the ant social system. In prep.
Aim of the study
The presence of a parasite in a system (Rickia wasmannii) could decisively
influence the infiltration success of another parasite (Maculinea spp) either
negatively or positively. During our study we investigated in laboratory conditions
the differences between R. wasmannii infested and non-infested Myrmica
scabrinodis colonies with regard to the adoption process (duration and complexity)
of caterpillars of four different Maculinea taxa: M. alcon ruiata, M. a.
peuoathe, M. nausithous kijevensis and M. teleius (Fig.2.2. a,b,c,d).
Materials & methods
The infested and non-infested Myrmica scabrinodis colonies were
collected (12.05-21.07.2012) from Fațul Domnesc, where all four Maculinea taxa
cohabit stopiall Tiuș et al. a) and parasitize Myrmica scabrinodis
(Tartally et al. 2008, pers obs.). Maculinea caterpillars of the fourth larval stage
were obtained by collecting host plants with larvae of the different species from
the 30th of June until the 15th of August (2012).
The adoption experiment was carried out from July to August 2012. In a
formicarium (queenless nests containing 50 workers Fig.2.2. e,f) a single Maculinea
caterpillar of the 4th larval instar was introduced. The time elapsed from the
placing of the caterpillar until its discovery by ants and its transportation in the
shelter (Fig.2.2.g), respectively, was recorded on a minute per minute basis for 120
minutes. In addition, the behavioural responses of ants towards the caterpillar
were also noted [antennation, ant harvesting the DNO secretions droplets
(Fig.2.2.h), transportation of a caterpillar in to the nest etc.]. Altogether 132
caterpillars of the four Maculinea forms were included in the experiment: 65
infested colonies versus 67 non-infested colonies.
The effect of infestation and of the effect of Maculinea species on the
adoption rate of caterpillars was analyzed with a Cox regression approach
(proportional hazard, efron method for handling ties). The sum of interactions
occurring between ant workers and caterpillar was analyzed with Generalized
Linear Mixed Model (GLMM) approach, Poisson error.

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Results & Discussion
Infestation of Myrmica scabrinodis ants with the fungus Rickia wasmannii
influences differentially the adoption success and the duration of the adoption
process of Maculinea species. M. alcon peuoathe (significant) and M.
teleius (marginally significant) caterpillars were adopted sooner and in higher
percentage by infested ants, whereas M. nausithous kijevensis (marginally
significant) by non-infested ants. In the case of M. alcon ruiata the differences
between infested and non-infested were not significant. The different effects of
fungal infestation of Myrmica scabrinodis ants could be the result of local
specialisation of Maculinea species for infested or non-infested ants located in the
vicinity of specific host plants.
In this study we discovered that Myrmica scabrinodis ants, irrespective of
the infestation, perform a specific adoption rituals towards Maculinea species. The
most simple and the shortest adoption process (seconds-5min) was in M. alcon
peuoathe (Fig.2.1). The longest (50 min) and complex (high and divers No
of behavioral interactions) adoption ritual was with M. teleius caterpillars. In M.
alcon ruiata the adoption process was longer (30 min) (Fig.2.1) and more
complex than in M. alo peuoathe. M. nausithous kijevensis caterpillars
were significantly more inspected (No of antennation) but the duration of adoption
process was shorter (15 min) than M. alcon ruiata and M. teleius.
Fig. 2.1. The duration af adoption process of Maculinea species with Myrmica
scabrinodis infested ants (red) and not infested (white).

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Fig.2.2. Photo: Natalia Timuş
a) Maculinea alcon “cruciata”. b) Maculinea alcon “pneumonanthe”. c) Maculinea teleius. d)
Maculinea nausithous kijevensis. e) Formicarium f) Myrmica scabrinodis colony wth 50
worker ants. g) Rickia wasmannii infested ant of Myrmica scabrinodis transports M. teleius
caterpillar. h) Myrmica scabrinodis ant “imbibe” the droplet secreted by M. teleius dorsal
nectary organ.

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Chapterl 3. Survival and development of Maculinea caterpillars
in Myrmica scabrinodis colonies infested with Rickia wasmannii
versus non-infested
Some results of this study in: Markó B., Csata E., Timuș N., Hughes M., Tartally A., Csősz S.,
Rózsa L.: A unique multispecies parasitic system worth protecting in ants. In prep.
Aim of the study
We analysed the survival (days) and growth (mg) of caterpillars of four
different Maculinea taxa (M. alcon ruiata, M. a. pneumonanthe M. teleius,
M. nausithous kijevensis) in Myrmica scabrinodis host-ant colonies infested with
ectoparasitic fungus Rickia wasmannii versus not-infested colonies. The main
objectives was to identify: the effects of fungal infestation of Myrmica scabrinodis
ants on survival and growth of Maculinea caterpillars in the same laboratory
conditions; integration level of Maculinea species in Myrmica scabrinodis colonies;
differences between Maculinea uko speies (M. alcon ruiata, M. a.
peuoathe) and predatory species (M. teleius, M. nausithous kijevensis).
Materials & methods
Infested and non-infested Myrmica scabrinodis colonies and specific host
plants as source of Maculinea caterpillars were collected from Faţul Does.
Altogether 158 caterpillars of the four Maculinea taxa were included in the
experiment: 79 in colonies with infested ants versus 79 in colonies with non-
infested ants. Survival and growth (body mass changes) of Maculinea caterpillars
were assessed at 10-day intervals. The experiment was carried out from 1 July
2012 to 26 July 2013. The effect of infestation on survival of Maculinea caterpillars
was analysed wiht Generalized Linear Mixed Model approach. Statistical analysis
Mann-Whitney, Wilcoxon (comparison of general growth pattern of Maculinea
caterpillars in infested and non-infested ant colonies), Kruskal-Wallis (comparison
of general growth pattern between Maculinea species) were realised with the
program PAST (Hammer et al. 2001). Bonferroni correction was used for each post
hoc comparison.
Results & Discussion
Survival rate and median survival time ee highe i the ukoo
Maculinea species and they were very reduced in predatory species (Fig.3.1.). M.
teleius caterpillars survived until 20 weeks and M. n kijevensis until 14 weeks. The
ukoo caterpillars of the M. alcon ecotypes survived until 11 months inside ant
nests. The fungal infestation of ants influences differentially the survival of
Maculinea species (Fig.3.1.). The results showed a reduced survival rate (Fig.3.2.)
and median survival time of M. n. kijevensis caterpillars in infested colonies. The

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infestation of ants have a positve effect on M. teleius demonstrated by higher
values of survival rate (Fig.3.2.) and body mass (is the only case were the
infestation have an effect on growth of caterpillars). In M. a. ruiata survival
rate and median survival time, although marginally significant, were higher in
infested colonies. For M. a. peuoathe the difference between infested and
non-infested ants, was insignificant.
All studied Maculinea taxa (exception M. a. ruiata ) grew rapidly in
the first 10 days (Fig.3.2., Fig.3.3.d), achieving a specific post-adoption weight. M.
a. ruiata caterpillars grew constantly till pupation (Fig.3.2). The specific post-
adoption weight of M. a. peuoathe remained stable from August till March-
April, when it was possible to distinguish two groups of caterpillars, faster and
slower growing caterpillars, which may indicate the presence of polymorphic
larvae. Differences in general growth pattern between M. alcon ecotypes it is likely
to be a mechanism for synchronizing their flight periods with the availability
(phenology) of specific host plants (Sielezniew & Stankiewicz 2007).
Irrespective of the infestation the Maculinea caterpillars had a specific
level of integration in ant colonies. M. a. peuoathe and M. a. ruiata
were extremely well integrated in host ant colonies (Fig.3.3.a), M. n. kijevensis are
well integrated (Fig.3.3.b), and M. teleius caterpillars were tolerated or ignored by
host ants. M. n. kijevensis shoed ukoo and predatory behaviour, and most
important aspect was the re-adoption of caterpillars after reintroduction in their
host ant colonies. Ol ukoo M. a. peuoathe and M. a. ruiata
caterpillars became infested by R. wasmanii after one month spent in infested
colonies (Fig.3.3.e,f). The infestation of Maculinea caterpillars had no detectable
effect. Only a single caterpillar of M. alo peuoathe pupated wich was
infested with Rickia wasmannii. Also, I found infested M. alo peuoathe
caterpillars in the field (June 2013, Faţul Does).
Fig. 3.1. The median survival time of
M. a. “pneumonanthe” in infested
colonies (api) vs non-infested (apn),
M. a. “cruciata” (aci vs acn), M. n.
kijevensis (nki vs nkn), M. teleius (ti
vs tn).

16
Fig. 3.2. Survival curve (interrupted line) and growth curve (continuous line) of Maculinea
caterpillars in infested Myrmica scabrinodis colonies (red) and non-infested colonies (black).
0
10
20
30
40
50
60
70
80
90
100
adoptate
20
40
60
80
100
120
140
vara
toamna
iarna
growth (mg) Survival curve (%)
M. teleius, 2012
0
10
20
30
40
50
60
70
80
90
100
adoptate
20
40
60
80
100
120
140
vara
toamna
iarna
growth (mg) Survival curve (%)
M.n.kijevensis, 2012
0
10
20
30
40
50
60
70
80
90
100
adoptate
30
60
90
120
150
180
210
240
270
300
330
vara
toamna
iarna
primavara
vara
growth (mg Survival curve (%)
M.a. "cruciata", 2012-2013
0
10
20
30
40
50
60
70
80
90
100
adoptate
30
60
90
120
150
180
210
240
270
300
330
vara
toamna
iarna
primavara
vara
growth (mg) Survival curve (%)
M. a. "pneumonanthe", 2012-2013

17
Fig.3.3. Photo: Natalia Timuş
a) Myrmica scabrinodis ant transorts M. alcon “pneumonanthe” caterpillar. b) Adopted
caterpillar of M. n. kijevensis (arrow) among Myrmica scabrinodis ant brood, beneath floral
foam (artificial nest). c) Pre-adoption fourth instar caterpillar of M. teleius (2,9 mg). d) Post-
adoption fourth instar caterpillar of M. teleius (11,7 mg) after 10 days spent in host ant
colony e) M. alcon “pneumonanthe” caterpillar infested with Rickia wasmanii. f) M. alcon
“cruciata” caterpillar infested with Rickia wasmanii.

18
Chapter 4. Demographic parameters of Maculinea teleius and
M. nausithous kijevensis from “Dealurile Clujului Est”
Parts are published as: Vodă et al. 2010, Timuş et al. 2011
Aim of the study
We applied the mark-release-recapture (MRR) method to study the adult
populations of the M. teleius and M. nausithous kijevensis species with the aim of
gathering knowledge of importance for their conservation. In the study area both
species have similar ecological requirements: Sanguisorba officinalis as host plant
and Myrmica scabrinodis as host ants.
Materials & methods
The population of Maculinea teleius (Fig.4.2a) from Fațul Does was
analysed with mark-release-recapture (MRR) method in 2009, 2010 and 2011. The
MRR method was applied in 2011 on syntopic populations of Maculinea teleius and
M. nausithous kijevensis from Faţul Does (Fig.4.2.a,c) and Faţul “ătes
(Fig.4.2.b,d). Data were analysed separately for each species with the Cormack-
Jolly-Seber type constrained models (Schwarz & Arnason 1996) using the program
MARK 6.0 package (Cooch & White 2010).
Results & Discussion
The adults of M. teleius and M. n. kijevensis flew for approximately 7
weeks, between the 7th July – 25th August. The average life span of both species
was 3-5 days (Table 4.1). The sex-ratio dynamics within season of M. teleius and M.
n. kijevensis always showed a pattern of proterandry, a common fenomena in
Maculinea butterflies (Nowicki et al. 2005a).
According to the estimates, the size of the population of M. teleius from
Fațul Does -2011) was 1000 – 1800 individuals and 1200 individuals
for M. n. kijevensis in 2011. In the year 2011 in Faţul “ătes a population of 2800
individuals was estimated for M. teleius and 1700 individuals for M. n. kijevensis
(Table 4.1, Fig.4.2. a,f). The M. teleius experienced small demographic fluctuations
through the years.
For both species within-season daily population size dynamics and the
recruitment of both males and females consistently followed a bimodal pattern
(Fig.4.1.), as was showed in other studies (Nowicki et al. 2005b).
Only 3 individuals of M. teleius and 2 individuals of M. n. kijevensis
changed sites. The results indicate that there are necessary measures (corridors
and stepping stones) to enhance metapopulation viability.

19
Tabelul 4.1. Basic parameters of populations of Maculinea teleius and M. nausithous
kijevensis (M.n.k.) as revealed by MRR studies in Fânațul Domnesc (FD) and Fânațul
Sătesc (FS).
N.M. – no of marked individuals, N.E. – no of estimated individuals, p -average daily capture
probability; φ-average daily survival rate; M - males, F -females, M : F- sex ratio (%).
Fig. 4.1. Within – season recruitment in the populations investigated: M. teleius and M. n.
kijevensis, Fânațul Domnesc, 2011.
0
100
200
300
400
500
600
700
individuals
M. teleius
M.n. kijevensis
Maculinea teleius
Study
area &
year
N.M.
N.E.
M : F
(%)
φ
p
Life span
M
F
M + F
M
F
FD 2009
268
1231
48 : 52
0,71
0,71
0,23
4,34
4,34
FD 2010
279
981
46 : 54
0,81
0,81
0,42
4,66
4,66
FD 2011
342
1774
55 : 45
0,81
0,75
0,20
4,73
3,54
FS 2011
332
2808
59 : 41
0,73
0,73
0,15
4,72
4,72
M.n.k.
FD 2011
235
1204
63 : 37
0,85
0,78
0,15
3,33
2,96
FS 2011
247
1676
60 : 40
0,81
0,59
0,20
4,85
1,93

20
Fig. 4.2. Photo: Natalia Timuş
Marked Maculinea teleius individuals in Fânaţul Domnesc a) and Fânaţul Sătesc b)
Marked M. nausithous kijevensis individuals in Fânaţul Domnesc c) and Fânaţul Sătesc d)
e) GPS coordinates of captured M. teleius butterflies in Fânaţul Domnesc and Fânaţul
Sătesc. f) GPS coordinates of captured M.n.kijevensis butterflies in Fânaţul Domnesc and
Fânaţul Sătesc (Luna de Jos, jud. Cluj).

21
Chapter 5. Mobility and spatial distribution within habitat of
Maculinea teleius and Maculinea nausithous kijevensis
butterflies
Results of this study in: Timuș N., Nowicki P., Rákosy L.: Within-population source-sink
dynamics in Maculinea butterflies. In prep.
Timuș N., Czekes Z., Craioveanu C., Nowicki P., Rákosy L.: Movement patterns of two
syntopic Maculinea species. In prep.
Aim of the study
Recent studies (e.g. Kőösi et al. , Whoff et al. , Noiki et al.
2013, Skórka et al. 2013a,b) revealed that M. teleius and M. nausithous species,
even occuring syntopically, have a regional and species-specific pattern of mobility
and spatial distribution, shaped by ecological requirements, arrangement of
resources within habitat, behaviour of resource exploitation (host plants and host
ants), physical structure of available habitats. The aim of our study was to indentify
the movement and spatial distribution patterns within habitat of syntopic species
M. teleius and M. nausithous kijevensis from Fațul Does. In the study site
both species have the same ecological requirements: Sanguisorba officinalis
utilised as host plant and Myrmica scabrinodis as host ants species.
Materials & methods
We used the mark-release-recapture (MRR) data from 2009, 2010 and
2011 and GPS coordinates. The mobility of butterflies was analysed with
Generalized Linear Mixed Model and Mann-Whitney U-test. The transition
probability of butterflies between high-quality habitat (source) and low-quality
habitat (sink) was calculated with multi – state models for live recaptures.
Results & Discussion
Our results showed that the mobility and spatial distribution of M. teleius
and M. nauisthous kijevensis fo Fațul Does have a specific pattern with no
significant inter-specific differences. The butterflies are characterised by home-
range behaviour and relatively low within-habitat mobility, the mean seasonal
flight distance was 100 – 200 m. The females of both species were more mobile
than males. The mobility of butterflies increased with season progression (Fig.5.1),
especially females tend to cover longer distances at the end of the flight period
(Fig.5.2, Fig.5.3), behaviour caused by oviposition preferences (Fig.5.5. a, b) and
host plants availability. The populations of M. teleius and M. nauisthous kijevensis
from Fațul Does are characterised by source-pseudosink dynamics within
habitat: source–high-quality habitat, consistent net exporter of organisms and
pseudosink - low-quality habitat, net importer, but without immigration they can
sustain populations and sometimes can even become netexporters (Watkinson &

22
Sutherland 1995, Boughton 1999). The transition probability of butterflies from
source to pseudosink increases rapidly after the peak of flight period (Fig.5.4,
Fig.5.5 e,f)). The tendency to leave the source was more pronounced in females
(Fig.5.4) than males, in order to avoid inter an intra-specific competition by finding
host plants with no eggs and in a suitable phenological stage.
Source-pseudosink dynamics of studied butterflies could be a regulating
factor of populations that mentain a diversity of syntopic Maculinea species in
Fațul Does, through absorbtion of the individuals excess by pseudosink,
thereby reducing the parasite pressure on Myrmica ants from source.
1JPZ s1
2 JPZ s1
0
60
120
180
240
300
360
420
480
540
distanta de zbor (m)
Fig. 5.2. The mean flight
distance of M. teleius
females in the first half (1JPZ
s1) and the second half
(2JPZ s1) of flight season,
MRR data from 2009- 2010
Fig. 5.1. The increasing of flight
distances of M. teleius and M. n.
kijevensis with season
progression (time in days
between the start of the season
and the middle of the period
between captures), MRR data
from 2009-2010, GLMM.

23
Fig. 5.4. – Probability (%) of dispersion from source to pseudosink of M. teleius and M. n.
kijevensis females, 2010 (Psi A to B, multi-state model).
1JPZ s1
2JPZ s1
0
100
200
300
400
500
600
700
800
900
distanta de zbor (m)
0.00
0.20
0.40
0.60
0.80
1.00
1.20
1.40
12
iulie
15
iulie
18
iulie
21
iulie
23
iulie
30
iulie
3
aug.
6
aug.
9
aug.
12
aug.
16
aug.
18
aug.
21
aug.
24
aug.
transition probability (%)
femele de M. n. kijevensis
femele de M. teleius
Fig. 5.3. The mean flight
distance of M. n. kijevensis
females in the first half
(1JPZ s1) and the second
half (2JPZ s1) of flight
season, MRR data from
2009- 2010

24
Fig.5.5. Photo: Natalia Timuş
a) Female of M. teleius (marked with number 54) ovipositing on young inflorescence of
Sanguisorba officinalis. b) Female of M. n. kijevensis (marked with number 645) ovipositing
on mature inflorescence of S. officinalis.
The maps shows the source (orange line) and pseudosink areas (green line) in Fânaţul
Domnesc.
Captured Maculinea teleius individuals in the first c) and second d) half of the fight season.
Captured Maculinea nausithous kijevensis individuals in the first e) and second f) half of
the fight season.

25
Chapterl 6. The land-use impacts on M. teleius populations
from Natura 2000 site “Dealurile Clujului Est”
Parts are published as: Timuş et al. 2011
Aim of the study
The aim was to identify in which type of land-use (abandoned, mown,
grazed) Maculinea teleius populations are the most prosperous . Another objective
was to find out if transect method could be useful to assess the land-use impacts
on M. teleius populations. According to our results obtained from MRR method,
transect counts, and information related to land-use history, we indicated the
most threatening factors for M. teleius populations in the study area, and
described specific conservation measures in order to insure the long term survival
of these butterflies.
Materials & methods
This study was carried out in three meso-higrophilous meadows: Fațul
Domnesc (41ha)- partially abandoned and intensively grazed by sheep (Fig.6.4.c),
Fațul “ătes ha - mown in a mosaic-like manner and partially grazed by
sheep (Fig.6.4.b), “eheliște ha – mown in a mosaic-like manner, the grazing is
forbbiden (Fig.6.4.a). In each meadows we delimited the habitats with Maculinea
teleius: 20 ha in Fațul Does FD,  ha Fațul “ătes F“,  ha “eheliște á
(SA) and , ha “eheliște B “B. We recorded M. teleius individuals with transect
method in FD, FS, SA and SB, and a mark-release-recapture (MRR) method was
applied in FD and FS. The MRR data were analysed using the program MARK 6.0
package (Cooch & White 2010). Transects data and MRR data were compared
through an analysis of variance (one-way ANOVA).
Results & Discussion
Using the transect method the lowest values were recorded i Fațul
Domnesc (53 individuals) and Fațul “ătes  individuals) (Fig.6.1., Fig.6.2.).
Highe alues ee otaied i “eheliște á  idiiduals) and “eheliște B (140
individuls) (Fig.6.1., Fig.6.2.). Transect count results are highly correlated with
estimates obtained with MRR (Fig.6.2.). The estimated populations of M. teleius
were: 5276 individuals i “eheliște B, 2808 de indivizi î Fațul “ătes,  de
idiizi î “eheliște á,  de idiizi î Fațul Does (Fig.6.3.).
M. teleius thrives in meadows (Dealurile Clujului Est) were mowing was
applied tardily, asynchronously and in a mosaic-like way. The grazing on these
medows was forbbiden. We recommend avoiding the mowing and grazing of
meadows on which M. teleius colonies were identified between 1.06 – 10.09. A
compromise solution, which could be convenient for farmers and hay production

26
and also maintaining M. teleius population at a balanced level, is that of mowing
being applied after 25.08. Transects method may be a useful tool in tracking
population trends over time and their response to changing land use.
The main reason for the decline or the extinction of some populations of
M. teleius in the investigated area is the conversion of grasslands to arable land
(2013, 2014) and the mowing of meadows in the flight season of these butterflies
(2012) (Fig.6.4.d). Factors that have a negative impact on the populations of
Maculinea and their habitat are: the cessation of traditional agricultural practices
(grazing and hand-mowing) and abandonment of land, construction plans,
drainage of humid areas (Fig.6.4.f) and the alteration of proper, benefic habitats
for Large Blues by planting Robinia pseudoacacia, Pinus sylvestris or Pinus nigra.
Usually, in spring and autumn, farmers are setting on fire the dry vegetation
(Fig.6.4.e) but the effects of this impact are yet unknown.
Fig. 6.1. Number of M. teleius individuals recorded with transect method and number of
individuals marked with MRR method, 2011.
Fig. 6. 2. Number of recorded M. teleius
individuals with trensect counts in Fânațul
Domnesc (FD), Fânațul Satesc (FS),
Secheliște A (SA), Secheliște B (SB),
2011.
Fig. 6.3. The estimated population size
(MARK) of M. teleius in Fânațul Domnesc
(FD), Fânațul Satesc (FS), Secheliște A
(SA), Secheliște B (SB), 2011.
0
10
20
30
40
50
60
70
80
9 iulie
11 iulie
13 iulie
15 iulie
18 iulie
20 iulie
27 iulie
3 aug.
5 aug.
7 aug.
12 aug.
15 aug.
18 aug.
22 aug.
Individuals
Fânațul Domnesc
0
10
20
30
40
50
60
70
80
90
100
15 iulie
19 iulie
28 iulie
4 aug.
6 aug.
8 aug.
12 aug.
15 aug.
18 aug.
22 aug.
Individuals
Fânațul Sătesc
marked MRR
transect
aband.
grazed
mown
mown
grazed
mown
0
50
100
150
FD (20
ha)
SA (15
ha)
FS (5
ha)
SB (3.8
ha)
individuals
aband.
grazed
mown
mown
grazed
mown
0
1000
2000
3000
4000
5000
6000
FD (20
ha)
SA (15
ha)
FS (5
ha)
SB (3.8
ha)
individuals

27
Fig.6.4. Foto: Natalia Timuş
a) Sechelişte – mown in a mosaic-like manner, the grazing is forbbiden. b) Fânaţul Sătesc
- mown in a mosaic-like manner and partially grazed by sheep. c) Fânaţul Domnesc-
abandoned and partially and intensively grazed by sheep. d) Site with syntopic Maculinea
species from Fânaţul Sătesc – mown in July 2012.
e) Burnt grass, Molinia caerulea patches in Fânaţul Domnesc (May 2011). f) Drainage
canal (2009) on site (southern exposed) opposed to the Maculinea site Fânaţul Domnesc
(northern exposed).

28
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Ioan Nemeş: 29-46.
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List of publications
Pu bli shed a rti c l es
Tiuș N., Craioveanu C., Sitaru C., Rus A., Rákosy L. (2013a) Differences in adult phenology,
demography, mobility and distribution in two syntopic ecotypes of Maculinea alcon
(cruciata vs. pneumonanthe) (Lepidoptera: Lycaenidae) from Transilvania (Romania).
Entomologica romanica 18: 21-30.
Tiuș N., Constantineanu R., Rákosy L. (2013b) Ichneumon balteatus (Hymenoptera:
Ichneumonidae) – a new parasitoid species of Maculinea alcon butterflies
(Lepidoptera: Lycaenidae). Entomologica Romanica 18: 31-35.
Tiuş N., Vodă ‘., Paulii I., Cişa á., Popa ‘., ‘kos L.  Maageetul pajiştilo
ezohigofile de pe Dealuile Clujului Est Tasilaia petu poteţia şi
conservarea speciei Maculinea teleius (Bergsträsser 1779) (Lepidoptera:
Lycaenidae). Volumul de luăi al “ipozioului Biodiesitatea şi Maageetul
Isetelo di ‘oia, “ueaa, -25 septembrie 2010, în memoria
etoologului uoiea Ioa Neeş: -46.
Vodă ‘., Tiuş N., Paulii I., Popa ‘., Mihali C., Cişa á., ‘kos L.  Deogaphic
parameters of two sympatric Maculinea species in a Romanian site (Lepidoptera:
Lycaenidae). Entomologica romanica 15: 25-32.
Avram A., Cîmpean M., Jurca A., Tiuş N. (2009) Water quality assessment using biotic
indices based on benthic macroinvertebrates in the Somesul Mic catchment area.
“tudia Uiesitatis Baeş-Bolyai, Biologia 1: 61-70.

33
Ma nus cri pts in pr epa rat ion :
Tiuș N., Csata E., Witek M., Baik H, Czekes )., Eős K., ‘kos L., Markó B.: Parasitic fungi
as key to the ant social system.
Markó B., Csata E., Tiuș N., Hughes M., Tatall á., Csősz “., ‘ózsa L.: A unique multispecies
parasitic system worth protecting in ants.
Tiuș N., Nowicki P., Rákosy L.: Within-population source-sink dynamics in Maculinea
butterflies
Tiuș N., Czekes Z., Craioveanu C., Nowicki P., Rákosy L.: Movement patterns of two
syntopic Maculinea species.
Pr ize s:
Kutter – Prize for the best poster, 3rd Central European IUSSI Meeting, 2013
In ter nat ion al c onf ere nce s:
Natalia Tiuş, Markó Bálint, László Rákosy (2013) Infestation of Myrmica scabrinodis with
Rickia wasmannii (Ascomycetes: Laboulbeniales) aids the infiltration of socially
parasitic Maculinea species (Lepidoptera: Lycaenidae) differentially. 5th Central
European Workshop of Myrmecology, Innsbruck, Austria.
Natalia Tiuş, Markó Bálint, László Rákosy (2013) Fungal infestation in Myrmica host ant
influences the adoption succes of Maculinea caterpillars. 3rd Central European IUSSI
Meeting, Cluj-Napoca, România.
Craioveanu Cristina, Tiuş Natalia, Sitar Cristian, László Rákosy (2011) Population dynamics
in Maculinea alcon and Mauliea xerophila in semi natural grasslands in
Transylvania, Romania. Laufen, Germania.
ádei Cișa, Cistia “ita, Natalia Tiuş, László Rákosy (2011) The fragmentation of
grassland habitats as a consequence of land use and land use abandonment. Ethnic
Landscapes & Ethno-Ecosystems – interdisciplinary workshop. Cluj-Napoca,
România.
Ige Paulii, “ai Bădăău, Cistia Maloș, Moia Beldea, ‘alua Vodă, Natalia Tiuș,
ádei Cișa, Lszló ‘kos  Vegetatio sue of the ha eados i the

34
poposed Natua  site Easte Hills of Cluj Taslaia, ‘oaia. th
European Dry Grassland Meeting. Smolenice, Slovacia.
László Rákosy, Raluca Voda, Natalia Tiuş, Ciprian Mihali (2010) Habitat management for
Maculinea in Romania. Laufen, Germania.
László Rákosy, Raluca Voda, Natalia Tiuş, Ciprian Mihali (2009) Maculinea genus in
Romania. 16th European Congress of Lepidopterology. Cluj-Napoca, România.
Na tio n a l c onf ere nce s
Natalia Tiuş, Zsolt Czekes, Cristian Sitar, Cristina Craioveanu, László Rákosy (2014)
Mobilitatea la fluturii Maculinea teleius și Maculinea nausithous kijevensis în
interiorul habitatului. Al XXIV-lea Simpozion SLR, Cluj-Napoca, România.
Natalia Tiuș, Bálint Markó, László Rákosy (2014) “upaiețuiea și dezoltaea laelo de
Maculinea (Lepidoptera, Lycaenidae) în coloniile de furnici Myrmica scabrinodis
(Hymenoptera, Formicidae) infestate cu Rickia wasmannii (Ascomycetes,
Laboulbeniales) versus neinfestate. Al XXIV-lea Simpozion SLR, Cluj-Napoca,
România.
Natalia Tiuș, Bálint Markó, László Rákosy ( Cu poate iflueța ifestaea fuiilo
gazdă Myrmica scabrinodis (Hymenoptera, Formicidae) cu ciuperca Rickia
wasmannii ásoetes, Laouleiales suesul de adopţie a laelo de
Maculinea (Lepidoptera, Lycaenidae). Simpozion BIOTA, Cluj-Napoca, România.
Natalia Tiuş, Cistia “ita, Cistia Caioeau, Lszló ‘kos  Capaa eologiă –
o posibilă aeiţae petu flutuii Maculinea? Al XXIII-lea Simpozion SLR, Cluj-
Napoca, România.
Natalia Tiuş & László Rákosy (2012) Complexe parazitare din furnicarele de Myrmica
scabrinodis Nylander, 1846 (Hymenoptera, Formicidae). Al XXII-lea Simpozion SLR,
Galaţi, România.
Natalia Tiuş, Cristian Sitar, Cristina Craioveanu, László Rákosy (2012) Dinamica
populaţioală multianuală la Maculinea (= Phengaris) teleius Bergsträsser, 1779
(Lepidoptera, Lycaenidae) pe Dealurile Clujului Est (Transilvania, România). Al XXII-
lea “ipozio “L‘, Galaţi, România.
Natalia Tiuş, Raluca Vodă, Cistia Caioeau, ádei Cişa, Cistia “ita, Daga
Schmidt, Silvia Griger, Alexandra Rus, László Rákosy (2011) Date preliminare
efeitoae la diaia populaţioală a speciei Maculinea alcon (Denis &
“hiffeülle,  Lepidoptea: Laeidae di Faţul Does Dealuile
Clujului Est). Al XXI-lea Simpozion SLR, Cluj-Napoca, România.

35
Natalia Tiuş, Raluca Vodă, ádei Cişa, Cistia “ita, Cistia Caioeau, “ilia Gige,
Lszló ‘kos  Diaia populaţioală la Maculinea teleius (Bergsträsser,
 şi M. nausithous kijevensis (Sheljuzhko, 1928) (Lepidoptera: Lycaenidae) din
Faţul Does Dealuile Clujului Est. ál XXI-lea Simpozion SLR, Cluj-Napoca,
România.
Natalia Tiuş, ‘alua Vodă, Cipia Mihali, ádei Cișa, Czekes Zsolt  Codui și
stategii seete î elația flutue Maculinea) - fuiă Myrmica). În cadrul
ofeițelo știițifie de la depataetul de Taooie și Eologie al Faultății de
Bilogie și Geologie UBB, Cluj-Napoca, România.
Natalia Tiuş, Raluca Vodă, Ige Paulii, ádei Cişa, ‘ăzvan Popa, László Rákosy (2010)
Maageetul haitatelo di peisajul ultual al Tasilaiei petu poteţia şi
conservarea genului Maculinea (Lepidoptera: Lycaenidae.).
Simpozion
Biodiesitatea şi aegeetul Isetelo di
‘oia, “ueaa, ‘oia.
‘alua Vodă, Natalia Tiuş, Ciprian Mihali, László Rákosy ( Ceetăi asupa uei
populaţii de Maculinea teleius Begstässe  şi M. nausithous ssp. kijevensis
(Sheljuzhko 1928) (Lepidoptera: Lycaenidae) din Transilvania. Al XX-lea Simpozion
SLR, Cluj-Napoca, România.
Natalia Tiuş, Raluca Vodă, Cipia Mihali, Lszló ‘kos  Mofologie lasiă şi
ofologie iohiiă la laele de Maculinea teleius Begstässe  şi M.
nausithous ssp. kijevensis (Sheljuzhko 1928).
Al XX-lea “ipozio al “oietății
Lepidopterologice Române (SLR), Cluj-Napoca, România.
Natalia Tiuş, ‘alua Vodă, Lszló ‘kos  Iteelația flutui și fuii î azul geului
Maculinea Lepidoptea. Î adul ofeițelo știițifie de la depataetul de
Taooie și Eologie al Faultății de Bilogie și Geologie UBB, Cluj-Napoca,
România.
‘alua Vodă, Natalia Tiuş, Ciprian Mihali, László Rákosy (2009) Date referitoare la
Maculinea teleius Berg., şi M. nausithous Berg. în România (Lepidoptera,
Lycaenidae). Al XIX-lea “ipozio al “oietății Lepidopteologie ‘oe, Galaţi,
România

36
Acknowledgements
The completion of this research would not have been possible without
the inestimable help of Prof. Dr. László Rákosy along with that of Octavian
Iarmenco. I am truly grateful to Prof. Dr. László Rákosy for his confidence in me
throughout the period I have carried out my PhD study. I am also grateful for all his
suggestions, guidance, and patience with which he has examined my work papers,
including the present study. My gratitude is also expressed for the boundless
understanding and support which Octavian Iarmenco has provided me throughout
these 5 years. It is largely due to his moral and financial support that this research
has been realizable.
I would like to extend my appreciation for the immense help of Cristina
Craioveanu, Bálint Markó and Piotr Nowicki. Their statistical analysis of data, ideas
and recommendations proved to be of considerable importance to this study. In
the same way, I convey my gratefulness to my colleagues Inge Paulini and Zsolt
Czekes.
I am also appreciative of all my colleagues which have participated in the
field works (Mark - Release - Recapture method): ‘alua Vodă, Cistia
Sitar, Cistia Caioeau, ádei Cişa, “ilia Giger, Vasilian Bojan, Dagmar
Shmidt, Alexandra Rus, Vlad Dincă. To Ige Paulii ad Eilia “toiao I ould like
to thank particularly for their field works and all the information related to the
land-use, as well as for the floristic data they have provided.
For the high-quality SEM images (scanning electron microscope) I very
much thank to my colleague Ciprian Mihali and Mr. Dr. Lucian Barbu Tudoran.
My sincere gratitude to Mr. Dr. Raoul Constantineanu for the
identification and preparation of Ichneumon balteatus individuals.
I would also like to express my gratitude to Valeria Chelaru for the English
proofreading.