UNIVERSITY OF AGRICULTURE SCIENCE AND VETERINARY MEDICINE CLUJ-NAPOCA FACULTY … · 2013-07-11 ·...
Transcript of UNIVERSITY OF AGRICULTURE SCIENCE AND VETERINARY MEDICINE CLUJ-NAPOCA FACULTY … · 2013-07-11 ·...
UNIVERSITY OF AGRICULTURE SCIENCE AND VETERINARY MEDICINE
CLUJ-NAPOCA
FACULTY OF HORTICULTURE
Biol. Ghizela-Daniela ALEC (VONICA)
[SUMMARY OF THE PHD THESIS]
IDENTIFYING AND EVALUATING ORNAMENTAL POTENTIAL OF SOME CENTAUREA
SPECIES FROM TRANSYLVANIA SPONTANEOUS FLORA WITH THE PURPOSE OF
COMPLIMENTING THE CULTIVATED ASSORTMENT
SCIENTIFIC COORDINATOR:
Prof. PhD. MARIA CANTOR
CLUJ-NAPOCA
2013
Ghizela D. ALEC (VONICA) Summary of PhD thesis
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THE PHD TABLE OF CONTENTS INTRODUCTION 10
Chapter 1
HISTORICAL RESEARCH OF CENTAUREA SPECIES WITH DECORATIVE POTENTIAL 12
1.1. Historical research of Centaurea species 12
1.1.1. Historical research in Romania 12
1.1.2. Historical research in Europa 17
1.2. Economical importance of Centaurea genus 25
1.2.1. The ornamental purposes 25
1.2.2. The medicinal purposes 30
1.2.3. Species of Centaurea genus as weed plants 31
1.2.4. Species of Centaurea genus with protective status 32
Chapter 2
INTERSPECIFIC AND INTRASPECIFIC DIVERSITY OF CENTAUREA GENUS FROM
TRANSYLVANIA
35
2.1. Systematic position of Centaurea species at national and international level 35
2.1.1. Centaurea subgenus 38
2.1.2. Cyanus subgenus 41
2.1.3. Lopholoma subgenus 42
2.2. Centaurea species from Transylvania 43
2.2.1. Morphological characteristics of Centaurea species from Transylvania 43
2.2.2. Chorology and ecological requirements of Centaurea species from Transylvania 61
Chapter 3
PHYSICAL AND GEOGRAPHICAL CONSIDERATIONS OF TRANSYLVANIA 71
3.1. Geographic location and limits 71
3.2. Geological characteristics 72
3.3. The relief of Transylvania 72
3.4. Hydrography 74
3.5. Climate 74
3.6. Soils 75
PERSONAL CONTRIBUTION
Chapter 4
OBJECTIVES OF RESEARCH 76
4.1. Morphological research objectives 76
4.2. Research objectives of acclimatization of Centaurea species with decorative potential 77
Chapter 5
RESEARCH MATERIAL AND METHODS 79
5.1. The materials and methods used in morphological research on Centaurea species 79
5.1.1. Biological materials used in research 79
5.1.2. Research methods used in the field 81
5.1.3. Research methods used in the laboratory 86
5.2. Materials and methods used in research acclimatization 90
5.2.1. Biological materials used in acclimatization 90
5.2.2. Methods used in the field research 91
5.2.2. Methods used in the laboratory research 92
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Chapter 6
RESULTS AND DISCUSSIONS 94
6.1. Morphological research in the Centaurea genus from Transylvania 94
6.1.1. Centaurea subgenus 94
6.1.2. Cyanus subgenus 175
6.1.3. Lopholoma subgenus 178
6.2. Research of morphological and ecological characters of Centaurea species with
decorative potential grown in the garden condition
193
6.2.1.Centaurea stenolepis subsp. ratezatensis (Prod.)(comb. nov.) 195
6.2.2. Centaurea orientalis L. 208
6.2.3. Centaurea atropurpurea Waldst. et Kit. 217
6.3. Key determination of Centaurea genus from Transylvania 231
Chapter 7
GENERAL CONCLUSIONS 237
REFERENCES 247
ALPHABETICAL INDEX OF AUTHORS 262
INDEX OF TABLES 268
INDEX OF FIGURES 273
INDEX OF PHOTOS 276
ANNEX 280
LIST OF SCIENTIFIC PAPER, WITH SUBJECTS FROM PHD THESIS 283
SUMMARY OF PHD THESIS IN ENGLISH AND ROMANIAN LANGUAGE 285
KEY WORDS
Centaurea Subgenus
Interspecific diversity
Morphology and morphometrie
Polymorphism
Polyploids
Hybridization
Systematic
Ordination analysis
Classification analysis
Canoco software
Past software
Acclimatization
Dichothomous key
Transylvania
Romania
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INTRODUCTION
Centaurea genus is considered from a systematic point of view, a critical group of
plants. This situation is mainly due to the result of the large number of the natural hybrids
between plant species, with high polymorphism. This polymorphism is the result of their
hybridization capacity or due to incomplete diagnoses or the lack of many species
description. For these reasons, the Centaurea genus has not been peer-reviewed for
decades and the newest studies took the results from the beginning of XX th
Century,
which come into conflict data with their geographical spread. This entire situation leads
to the Centaurea genus „marginalization” and failure to address in different scientific
works.
Until now, the unresolved taxonomic problems plus the fact that Centaurea
species are plants with tall steams, not easy to collect and keep it in the herbarium, with
common insect attacks, lead the plant researcher to avoid them.
Many scientific papers occurred comments like: „can not correctly determine
the...”, „ no one knows what species/ hybrid is this...”, „the species have an uncertain
taxonomic status...” etc.
The approach of Centaurea themes, has gone from the vacuum information, the
only taxonomic study of the groups is due to the studies developed by Juliu Prodan and
Erasmus Nyárády, known both in Romania and Europa (PRODAN & NYÁRÁDY, 1964). It
is too difficult to fill the gap of Nyárády and Prodan studies, but the research from this
thesis make a contribution to clarify taxonomic issues and to stimulate the research on
this group. On the other hand, the knapweeds are very attractive plants for decorating the
parks, gardens, balconies or for cat flower arrangements and bouquets.
Thus, the motivation to study these plants is due to the ornamental value given by
the variety of colors, robustness and their beauty. On the other hand, the chance to
elucidate the taxonomic problems, rewarding professional and systematic studies of the
group increases.
Due to the large number of Centaurea species at the national level, the research
was reduced to the Transylvanian level, which is an interesting area from a climatic and
relief forms point of view. This area is an inner-mountain shelter for diversity of
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vegetation and flora and here are found many species of Centaurea but also the rare
species from the flora of Romania and Europe.
During the research field, the human impact is often felt and for this reason is
necessary to protect and preserve the rare species of Centaurea.
This thesis is divided into seven chapters, it contains a number of 318 pages which
include 75 tables, 45 figures or graphs and 40 photo plates. The first three chapters have
an informational and general character and the last chapters bring personal contribution.
Chapter 6, Results and discussions, is divided into three subsections and the first section
tackles the taxonomic and morphological problems of Centaurea genus from
Transylvania. The second section represents the data recorded after the acclimatization of
Centaurea species to the garden condition and the dichotomous key of Centaurea genus
from Transylvania is the third subsection of this chapter.
The documentation for the correct understanding of the topic taken under study,
the research organization, the carriage of the observations and measurements, the
statistical processing, the interpretation and presentation of the results and, finally the
development of the thesis consist in the consultation of 205 references. In addition to the
references, the thesis also includes the alphabetical index of authors, tables’ index, graphs
and photo plate’s index and annex. Preliminary results of research over the years are
materialized in 11 scientifically papers presented and published in national and
international symposiums (see the list of scientific papers with thesis subject who is
added at the end of the thesis).
CHAPTER 1
THE HYSTORICAL RESEARCH OF CENTAUREASPECIES
1.1. HISTORICAL RESEARCH OF CENTAUREA SPECIES FROM ROMANIA AND EUROPE
Studies of Centaurea genus began with the first appearance of floristic research,
made by various botanists in Transylvania. The most important research of the genus, at
the national level, belongs to Professor Dr. Julius Prodan, from the Academy of
Agriculture Cluj (today UASVM Cluj-Napoca). He described in the first monograph of
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Centaurea genus (1930), many taxons and hybrids form Cluj surrounding and Dobrogea
area. This monograph contains 168 taxa identified in Romania, including hybrids. For a
long time, this monograph which was the result of Prodan’s work has been a major
resource (the only way) for botanists, because in describing of the species, are graphically
detailed the inflorescence items which are very important issues in species recognition.
Besides the works of Prodan, the Nyárády's studies have a great importance in the
evolution of the Centaurea genus study at national and Transilvanian level. Nyárády E. I.
tried to solve the "mystery" of this kind of morphological variability, describing many
taxa and hybrids of Centaurea species (NYÁRÁDY, 1943(a), 1943(b)).
Currently, the floristic research made on the Centaurea genus, focused on
determining species in Romania and Transylvania Plateau and was based on information
provided by the foreign papers, thus raising the quality of the information related to the
Centaurea genus issue. These works draw attention to the morphological variability and
hybridization of these species, indicating that the information provided by morphological
determination and separation are often insufficient and research must be completed with
molecular and chorological information. Taken these aspects into consideration, several
species of Romanian flora were put in synonymy or were misidentified (CIOCÂRLAN,
2009). Nevertheless, the studies of Ciocârlan (2009), compared with Prodan’s research
(1930), reduce the number of certain species on the Romania flora, from a few hundred to
54 species and he considers the presence of introduced species as doubtful or wrong in
Romanian flora (CIOCÂRLAN, 2008, 2009).
Regarding the systematic species of Centaurea genus, note that the last approach
of this kind of research was made in Romania by Julius Prodan, at 1930th
and it was based
on the Augustus von Hayek view (1901), which was a less conservative approach,
dividing the genus into seven genera.
In Europe, the Centaurea genus has been extensively studied, proved by numerous
studies on the systemic group, it updated the name of the species and the geographic
distribution of Centaurea species (BENTHAM &HOOKER, 1873; BREMER, 2004;
CASSINI,1829; FLANN et all., 2010; GREUTER, 2006–2009; KOUTEKÝ et all., 2012. PÉTIT
et all., 2001; TUTINet all., 1964-1980; EURO+MED, 2006; Flora Europaea -http://rbg-
web2.rbge.org.uk). Likewise, the taxa with uncertain positions have been the subject of
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thorough studies from genetically and palynologically point of view, which relieved the
positioning decision in phylogenetic system.
In Europe, the most important studies have adopted the conservationist
classification of the Centauries genus, bringing the seven types of Bentham and Hooker
(1873) in three subgenera (Centauries s. str., Cyanus, Lopholoma), boundary
classification which has been confirmed by molecular-palynological studies and
biogeographically patterns (FONT, 2007; GARCIA-JACAS et all., 2001; SUSANNA et all.,
1995; WAGENITZ & HELLWIG, 1996).
On the other hand, the number of species in the entire genus has been constantly
changing, depending on the evolution of genetic and morphological studies. Therefore,
the Centaurea species of the old subgenus has now become separate genera, as
Rhaponticum and Phesellus, and based on genetic analysis, the species C. ruthenica and
C. marschalliana were excluded from the current classification and included the Cnicus
benedictus species (AGABABIAN, 1997; ŐZLERet all., 2009).
1.2. THE ECONOMIC IMPORTANCE OF THE CENTAUREA GENUS
Centaurea genus has caught the attention of specialists due to the ornamental
quality and healing importance. At the national level, Centaurea species were introduced
only one or two native species in the culture as ornamental species, the rest of species
being exotic which have adapted more easily to climatic and soil conditions.
Floriculture was included in courses, the descriptions of just a few decorative
Centaurea species: C. moschata, C. cyanus, C. dealbata, which are imported from abroad
(BUTA et all., 2009; CANTOR & ZAHARIA, 2002; CANTOR &POP, 2005). Therefore, the
flower crops, parks, private gardens and pots have been lacked by the presence of native
flora, the beautiful "cornflowers".
As medicinal plant, C. cyanus L. (Bachelor's button) is well known for diuretic,
astringent, stomachic, hypoglycemic, eye anti-inflammatory and relieving properties
(ALEXAN et all., 1988; COICIU & RÁCZ, 1962; DRĂGULESCU, 1995, 2008; PÎRVU et all.,
2008).
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Attention of the agronomy specialists turned on the C. cyanus species when it
became a threat to cereal crops, acting in this way, but also leading to reduced
populations of inventory floristic (CIOCÂRLAN, 2004).
In terms of the protective status of Centaurea species, in Romania only two
Centaurea taxa are threatened with extinction at the Europe level, according to the
protected plant list of the Appendix II b from the Habitats Directive (92/43/EEC).At
Transylvania level, the protective species status is including three endemic Centaurea: C.
phrygia L. ssp. rarauensis (Prod.) Dostál, ssp. ratezatensis (Prod.) Dostál and C.
pinnatifida Schur (DIHORU & PÂRVU, 1987; DIHORU & NEGREAN, 2009; DOSTÁL, 1976;
SÂRBU et all., 2007).
CHAPTER 2
INTERSPECIFIC AND INTRASPECIFIC DIVERSITY OF
CENTAUREAGENUS FROM TRANSYLVANIA
Centaurea Lineé genus is part from Asteraceae (Compositeae) family, Cardueae
tribe, Centaurineae subtribe. Regarding the systematic analysis under genus level is
corroborated by morphological, palynological, genetic studies and geographical patterns
(SUSANNA & GARCIA-JACAS, 2007).
Currently, in Romania, it does not exist an updated classification system, to take
account of recent molecular or biogeographically studies, therefore this thesis will adopt
the systematic classification proposed by Garcia-Jacas and Susanna (1992), which is
largely over lapping with the classification of Wagenitz and Hellwig (1996) (Tab. 2.1).
The identified species from Transylvania are systematically assigned to three subgenera
(Centaurea s. str., Cyanus, Lopholoma) and eigth sections (Acrolophus, Jacea,
Lepteranthus, Calcitrapa, Cnicus, Cyanus, Protocyanus, Acrocentron).
In Centaurea s. str. subgenus were identified in Transylvania the following
species: C. stoebe, C. micranthos, C. diffusa, C. reichenbachii, C. triniifolia, C. jacea
with subspecies banatica and angustifolia, C. nigrescens, C. phrygia with subspecies
phrygia, erdneri, pseudophrygia, indurata, ratezatensis, carpatica, rarauensis,
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Table 2.1.
Systematic classification of Centaurea species from Transylvania*
Subgen
Subgenus
Secţie
Section
Subsecţie
Subsection
Specie
Species
Centurea s.str.
Acrolophus
(Acrolophus,
Willkommia,
Phalolepis)
Maculosae
C. stoebe
subsp. stoebe
subsp. micranthos
C. reichenbachii
C. triniifolia
Cylindraceae C. diffusa
Jacea Jacea (Juss.) Koch
C. jacea
subsp. angustifolia
subsp. bannatica
Nigrescentes, C. nigrescens
Lepteranthus (DC.)
DC Phrygia
C. phrygia
subsp. phrygia
subsp. erdneri
subsp. pseudophrygia
subsp. indurata
subsp. carpatica
subsp. ratezatensis
subsp. rarauensis
C. stenolepis
subsp. razgradensis
subsp. stenolepis
C. uniflora subsp. nervosa
C. macroptilon subsp.
oxylepis
Calcitrapa
Eucalcitrapa
C. calcitrapa
C. iberica
Mesocentron C. solstitialis
Cnicus - C. benedicta
Cyanus
Cyanus - C. cyanus
Protocyanus
C. triumfetti
subsp. axillaris
subsp. triumfetti
subsp. adscendens
subsp. stricta
subsp. pinnatifida
C. mollis subsp. mollis
Lopholoma Acrocentron
Acrocentron C. atropurpurea
Lopholoma
C. orientalis
C. kotschyana
C. scabiosa
subsp. spinulosa
*după / after: Garcia-Jacas & Susanna, 1992; Wagenitz & Hellwig, 1996
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C. stenolepis, C.macroptilon subspecies oxylepis, C. uniflora subspecies nervosa, C.
calcitrapa, C.
sostitialis, C. iberica, C. benedicta. The species C. cyanus, C. mollis, C. triumfetti with
subspecies axillaris, stricta, adscendens, pinnatifida mentionated in Transylvanian flora,
are part from Cyanus subgenus and C. orientalis, C. atropurpurea, C. kotschyana and C.
scabiosa with subspecies spinulosa are reprezenatives for Acrocentron subgenus.
CHAPTER 3
PHYSICAL AND GEOGRAPHICAL CONSIDERATIONS ON
INTRA-CARPATHIAN AREA STUDIED (TRANSYLVANIA)
Intra-Carpathian area of Romania, where the morphological studies of the
Centaurea genus were taken, overlaps with historical and geographical area of
Transylvania.
The depression hillside of Transylvania is central physical-geographical unit to
our country and it is limited by mountainous slopes of the three mountain branches: the
Eastern Carpathians, Southern Carpathians and the Western Carpathians with Apuseni
Mountains and intra-Carpathian yoke.
CHAPTER 4
RESEARCH OBJECTIVES
To achieve the goal regarding the identification of Centaurea species with the
decorative value from spontaneous flora of Transylvania and with the purpose of
complimenting the cultivated assortment, objectives were divided into two parts: the
objective of identifying morphological diversity and systematic of Centaurea species and
objectives of acclimatization Centaurea species with decorative value in the garden
conditions.
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CHAPTER 5
MATERIALS AND METHODS OF RESEARCH
In order to solve the problems related to taxonomy, detailed studies were made on
the polymorphic species that are relatively easy crossing, giving a rise number of
polymorphic individuals. In the wild flora of Transylvania were identified the following
polymorphic Centaurea species that were tracked in-situ: C. stoebe, C. jacea, C. phrygia
and the other species that are part of Phrygia group (C. phrygia subsp. indurata, C.
phrygia subsp. pseudophrygia, C. phrygia subsp. ratezatensis, C. stenolepis and C.
uniflora subsp. nervosa). Besides these polymorphic species, there were monitored the
following Centaurea species with decorative potential, in the wild flora of Transylvania:
C. cyanus, C. orientalis, C. atropurpurea and C. mollis.
To achieve the goals of acclimatization objectives of Centaurea species with
ornamental value, there were monitored in the garden condition the following species: C.
phrygia subsp. ratezatensis (C. stenolepis subsp. ratezatensis comb. nova), C. orientalis
and C. atropurpurea.
In addition to the biological material studied, for observations and measurements
on the field and laboratory, it was used an electronic caliper (Powerfix), USB Digital
microscope (400X zoom), Optical microscope B4 Optec, stereo magnifier Novex (40X)
and the electronic caliper.
For each species were made several morphological measurements (quantitative,
qualitative and reports) and the measurements were made on almost every vegetative
organ (stem, leaf, inflorescence, involucre bracts and achene). The numbers of characters
differ and depend on morphological peculiarity of each species.
As methods for the interpretation of morphological differences which occur in
some polymorphic species, we used ordination and classification statistical methods
computed by PAST-Paleontological Statistics 2.15 package (HAMMER, 1999-2012) and
CANOCO for Windows 4.5 package (TEER BRAAK & ŠMILAUER, 2002).
The ordination and classification analysis applied on theCentaurea genus are new
approaches in Romania, which are intended to clarify the morphological differences.
Morphological studies on the Centaurea genus which formed the basis of the thesis
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analysis were conducted in 2010-2012, on the spontaneous flora of Transylvania or on
the herbarium material (Herbarium of the Natural History Museum Sibiu-HSIB,
Herbarium of Cluj-Napoca Botanical Garden-HCL).
CHAPTER 6
RESULTS AND DISCUTIONS
In order to achieve the purpose and objectives of the research there were identified
in the Transylvanian flora a total of 22 species and 11 subspecies, based on the literature,
herbarium material (HSIB and HCL) and field research. For each subspecies there were
mentioned the synonymy and the herbarium material where reviewed from taxonomical
point of view.
Regarding the morphological monitoring in the spontaneous flora, it was made on
the polymorphic species that exhibited a high difficulty in determining species but also
have decorative values (VONICA & CANTOR, 2011(a), 2011(b)). For each species,
cytotypes or hybrids, discriminating morphological characters were presented, which can
differentiate the taxa and distribution maps on Transylvania area.
1. Regarding the differentiation of C. stoebe species (synonym with C. rhenana),
although it was known that the species has two ploidy levels (2x, 4x), we do not know if
in Transylvania flora could be found and how to differentiate the two cytotypes.
Following the ordination (PCA, CCA, RDA and UPGMA) and classification
analysis two cytotypes were found from morphologically point of view, which have been
identified in Europe flora by other botanists (Fig. 6.3).
C. stoebe subsp. stoebe (2n = 18), the diploid cytotype differs from the tetraploid
cytotype, C. stoebe subsp. micranthos (2n = 36), by inflorescence characters (length and
width-IL, IW), leaflets bracts and life cycle.
C. stoebe subsp. stoebe is, an annual species that grows well in open spaces, lack
of competition, the inflorescence height (measured from the base of the inflorescence to
the top of the last row of leaflets bracts) of 13.5 to 17 mm, appendage of leaflets bracts
has 6-10 pairs of fimbriae, lighter to the top and achene pappus is approximately half the
length of the achene.
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Figure 6.3. Discriminant score of two groups of C. stoebe individuals
(F 1,95; p 0,32)
C. stoebe subsp. micranthos is, a perennial species that grows in ruderalised
vegetation, with height from 19 to 23 mm of a cylindrical inflorescences, the appendage
of middle bracts with 4-7 pairs of fimbriae that have the same color throughout their
length and the pappus is about a quarter of the length of achene.
2. Regarding the differentiation of C. jacea individuals (other polymorphic group),
the results were obtained between this species and its hybrids with species from
Lepteranthus section. The ordination and classification analysis revealed four
discriminate morphological characters: appendage of middle bracts width (MW), the ratio
between length and width of leaflet bracts (BLW), leaf width (LW) and number of
fimbriae (FN) (Fig. 6.8).
C. jacea species differs, at first appearance by the hybrids, by shape of middle
bracts appendage, which is membranous, oval-rounded to C. jacea (5 to 5.6 mm) with
lacerate margin and lacking fimbriae, and by leaf width (6-8 mm in C. jacea). There
resulted hybrids from the crossing of C. jacea species and species of Lepteranthus
section differ from C. jacea by the irregular fimbriae on the margin which are focused in
small bouquets and by shape of middle bracts appendage to ovate-triangular to triangular-
lanceolate, with 3-4 mm appendage width, depending on the parent plant (Fig. 6.9). The
leaf width of hybrids achieves double the size comparing with C. jacea. Regarding the
-40 -32 -24 -16 -8 0 8 16 24 32
Scorul discriminant / Discriminant score
0
0,4
0,8
1,2
1,6
2
2,4
2,8
3,2
3,6
4
Fre
cven
ta /
Fre
quen
cy
Group I Group II
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ratio of the length and width of leaflet bracts it can be said that the C. jacea has 3.7-4.4
mm and hybrids 2.2-3.2 mm values.
Figure 6.8. CCA analysis of discriminant variable to
C. jacea species and its hybrids
-6 -4 -2 0 2 4 6 8
Axa / Axis 1
-4-2
02
46
8
Axa
/ A
xis
2
SAMPLES
jacea xjacea mel
C. jacea
C. x jaceaC. x melanocalathia
C. x melanocalathia
Figure 6.9. CCA plot of analysed individuals
(C. jacea, C. jacea hybrids and C. x melanocalathia)
Legend
jacea - C. jacea
xjacea –hybrid of C. jacea
mel - C. x melanocalathia
discriminant characters IL,
FN, LW, MW, BLW-
-10 -5 0 5 10 15
Axa / Axis 1
-15
-10
-50
510
15
Axa
/ A
xis
2
jacea
xjacea
mel
LW
IL
BLW
MW
FN
SPECIES ENV. VARIABLES
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3. Regarding the C. phrygia species (synonym with C. austriaca) it was found that
on flora of Transylvania two cytotypes (2x, 4x) are present, identified in Europe also as
C. phrygia subsp. phrygia (2n = 22) and C. phrygia subsp. erdneri (2n = 44) and
confirmed by the studies of this thesis (KOUTEKÝ et all., 2012). The two ploidy levels are
distinguished by morphological characteristics of middle bracts appendage and shape of
leaf. The middle bracts appendage of diploid cytotype has a width of 1 to 1.4 mm
compared with tetraploid cytotype (1-1.8 mm), the ratio between the length and width of
the leaflet bracts is 6.5 to the diploid or 5.8 to the tetraploid cytotype. Appendage of the
middle bracts has dark brown color at the base and light brown to yellow color to the
apex for diploid and dark brown to blackish brown at the base and dark brown at the apex
for the tetraploid. In terms of the leaf shape it can be said that the diploid cytotype has the
whole leaf slightly serrated on the margin with regular teeth and less than 1 mm long and
the tetraploid cytotype has lobed or toothed leaves with unequal lobes / teeth, larger than
1 mm long.In terms of geographical distribution it was found that C. phrygia subsp.
erdneri has a wider distribution in the studied intra-Carpathian area and it is present in
pastures where human influence felt, degraded grassland, grazed intensively etc. The C.
phrygia subsp. phrygia is distributed in the eastern part of Transylvania, in the meadows
of the Carpathians Mountains, depriving the southern, western and central Transylvania
(Photo 6.10).
Photo 6.10. Distribution of C. phrygia and C. erdneri taxa in Transylvania
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-9 -6 -3 0 3 6 9 12 15 18
Forta discriminanta / Discriminant
0
2
4
6
8
10
12
14
16
18
Frec
vent
a / F
requ
ency
4. Regarding the issue of the taxonomic approach of C. phrygia subsp.
ratezatensis, a diploid species (synonym with C. ratezatensis), it is observed that there is
a resemblance to the species C. phrygia subsp. pseudophrygia and C. stenolepis
(CIOCÂRLAN, 2009; DRĂGHIA et all., 2010). The ordination and classification analyses
revealed a morphological overlap of endemic species, C. ratezatensis, across species C.
stenolepis. Discriminating morphological characters evidenced by CCA are leaf width
(LW), branched stem length (measured from the first branching to the terminal
inflorescence-SBL), visibility of middle bracts appendage (MV) and the number of
fimbriae (FN) (Fig. 6.15).
Figure 6.15. CCA plots of C. stenolepis and C. ratezatensis species
Although, there are some characters that distinguish the two species, the
MANOVA test used to discriminating analysis (Hotelling) calculated a rate of
classification for both species by 92.05%, indicating that some of the morphological
characters of C. ratezatensis individuals overlap the morphological characters of C.
stenolepis (Fig. 6.16).
Figure 6.16. Cannonical score histogram of C. stenolepis and C. ratezatensis species
Legend
C. ratezatensis
C. stenolepis
-3.0 0.0 3.0
Axa / Axis 1
-0.6
0.0
1.0
Axa
/ A
xis
2
C. stenolepis
C. ratezatensis
SBLLW FN MV SPECIES ENV. VARIABLES
Ghizela D. ALEC (VONICA) Summary of PhD thesis
17
The classification analysis results confirmed that the C. ratezatensis taxon must be
included in the C. stenolepis species and treated as its subtaxons-Centaurea stenolepis
subsp. ratezatensis (Prod.) comb. nova (VONICA et all., 2013(a)).
5. Regarding the nomenclature approach of C. x prodanii and C. x devensis
hybrids, it was observed that the two hybrids are morphologically undifferentiated under
CCA analysis (VONICA et all., 2013 (b)). Classification analysis computed a score by
61.11% and morphological differences between the two hybrids are insignificant
according to the similarity test Hotelling's T–squared. (Fig. 6.21).
-0,9 -0,6 -0,3 0 0,3 0,6 0,9 1,2 1,5 1,8
Forta discriminanta / Discriminant
0
0,4
0,8
1,2
1,6
2
2,4
2,8
3,2
3,6
4
Fre
cventa
/ F
requency
Figure 6.21. Linear Discriminant analysis (LDA) of the
C. x prodanii şi C. x devensis hybrids
Due to lack of differential characters between the two hybrids, highlighted by
classification and ordination analyses, C. x prodanii described by Wagner (1910) as a
hybrid between C. phrygia and C. indurata and C. x devensis also described by Wagner
(1910) as a hybrid between C. stenolepis and C. phrygia are synonyms. The accepted
name for C. x prodanii was appointed by analyzing the geographical distribution of the
two parental species and the hybrid C. x devensis passed into synonymy.
6. Regarding the issue of taxonomic approach to species C. indurata (synonym
with C. phrygia subsp. indurata) may indicate that this species differs from C. stenolepis
with different ploidy level, length of leaf and inflorescence length. Although there is a
Legend
C. x prodanii
C. x devensis
Ghizela D. ALEC (VONICA) Summary of PhD thesis
18
partial morphological overlap and C. indurata species must not be treated as an inferior
taxon of the C. stenolepis species and, it will remain in the group Phrygia because it
requires additional morphological studies.
7. Regarding the morphological differences occurring in C. atropurpurea species
(subgenus Lopholoma), it can be said that they are directly related to the amount of
calcite in the soil. (VONICA et all., 2012). C. atropurpurea was monitored in several
populations of Transylvania and it was observed that in some populations (Turda Gorge -
CJ), the individuals of C. atropurpurea differ from higher values of lacinia length and
width, number of inflorescences and number of leaves compared to other individuals
populations (Lomb Mountain -CJ) (Fig.6.34). C. atropurpurea species is known as
calciphile plants and the higher values of morphological discriminating characters were
observed also in other populations.
Figure 6.32. PCA plots of C. atropurpurea species
(Jolliffe cut-off=0,74, confidence interval=95%)
8. Regarding the monitoring of rare species of Centaurea (C. stenolepis subsp.
ratezatensis, C. orientalis, C. atropurpurea) in the garden condition at Natural History
Museum Sibiu garden, it can be said the following (VONICA & CANTOR, 2012):
Componenta 1 (55,96 %, eigenvalue 6,53)
Co
mp
on
enta
2 (
18
,59 %
, ei
gen
val
ue
2,1
7)
Legend: Cheile Turzii
Dealul Lomb
Roşia de Secaş
Guşteriţa
Ghizela D. ALEC (VONICA) Summary of PhD thesis
19
-In terms of germination, it can be said that the greatest success recorded at C.
stenolepis subsp. stenolepis species (45.85%), followed by C. atropurpurea (18.66 %)
and C. orientalis (12.14 %).
- In terms of phenological phases of development, the C. atropurpurea species
blooming after 90-100 days from seeding followed by C. orientalis and later by C.
stenolepis subsp. ratezatensis.
- Between analyzed individuals developed in the garden conditions and
spontaneous flora, the ordination analyses showed a few discrimination morphological
characters. The plants of C. stenolepis subsp. ratezatensis developed in the garden
condition are different from plants from spontaneous flora, from the ratio between length
and width of the inflorescence, the ratio between length and width of the leaves and stem
height (Fig. 6.38).
Figure 6.38. CCA plots of two analysed groups (developed in the garden condition and
spontaneous flora) by C. stenolepis subsp. ratezatensis species
Individuals of the C. atropurpurea species developed in the garden condition
differ from individuals of spontaneous flora, in inflorescence and leaf width and length,
and between individuals grown in the garden and the spontaneous flora of the C.
orientalis species there is no difference (Fig. 6.43).
-2.0 0.0 4.0
Axa / Axis 1
-1.0
1.0
Axa /
Axis
2 Grãdinã
Flora spontanã
SLILW
LLW
Legend
ILW, LLW, SL –
discriminant
characters
Ghizela D. ALEC (VONICA) Summary of PhD thesis
20
Figure 6.43. CCA plot of two analysed group of C. atropurpurea species
(garden and spontaneous flora)
Recommendations:
C.stenolepis subsp. ratezatensis species grown in the garden conditions, along
with the other grassy species carpet, in shady and moist areas, it is recommended for
decorating brick walls or unsightly places in the surroundings of the house. It can be
recommended for floral spots in the parks or public gardens but can be also included in
the composition of flower carpet because it supports mowing grass and it has the ability
to regenerate from buds ingrown. The introduction of this species in the range of
ornamental species is a welcome ex-situ conservation of the species which is considered
endemic in Romania.
C. orientalis species is considered a relict species in Transylvania, but it is more
often outside the Carpathian arch and it is decorative in its yellow inflorescences. The
species is recommended for floral spots, intersections, parks and public gardens, in areas
with high insolation. It can be used as decorative species because the flowering stem is
straight and long, ended with one or two-three inflorescences. Being a perennial,
flowering in the first year, the vegetative success is very low and in unfavorable season
(winter), many plants do not survive at low temperatures and thermal protective measures
are required.
C. atropurpurea species grows well in the garden conditions, especially if the soil
is loose and contains calcite. The species can be used in the floral borders of the
intersections, or in parks and gardens. As a calciphile species, it can be recommended for
rocky gardens but with the condition to be limestone rocks. The species is also
-5.0 -4.0 -3.0 -2.0 -1.0 0.0 1.0 2.0 3.0
Axa / Axis 1
-1.0
-0.5
0.0
0.5
1.0
1.5
2.0
Axa
/ A
xis
2
Flora spontanã
Grãdinã
LWIL
Legend
IL, LW- discriminant
characters
Ghizela D. ALEC (VONICA) Summary of PhD thesis
21
recommended as cut flowers in floral arrangements and even as dried plant because the
inflorescence has a lacy appearance.
SELECTED REFERENCES
1. ALEXAN, M., O. BOJOR, F. CRĂCIUN, 1988. Flora medicinală a României, Ed.
Ceres, Bucureşti.
2. BENTHAM, G., J.D. HOOKER, 1873. Genera plantarum: ad exemplaria imprimis in
Herberiis Kewensibus servata definita, Ed. A. Black, 1862-
1883.http://dx.doi.org/10.5962/bhl.title.747.
3. BREMER, K., 1994. Asteraceae: Cladistics and Clasifications, Timber Press.
Portland.
4. BUTA, ERZSEBET, MARIA CANTOR, D. ZAHARIA, ADELINA DUMITRAŞ, A. ZAHARIA,
GEORGETA SZABO, M. BUTA, 2009. Description of some spontaneous species and
the possibilities of use them in the rocky gardens, J. Plant Develop., Iaşi, 16, 81-86.
5. CASSINI, H., 1829. Tableau synoptique des synanthéreés, În: Garcia-Jacas, Núria, T.
Uysal, K. Romashchenko, V. N. Suarez-Santiago, K. Ertuğrul, A. Susanna, 2006,
Centaurea Revisited: A Molecular Survey of the Jacea Group, Annals of Botany,
Oxford Journals, 98, 741–753.
6. CANTOR, MARIA, D. ZAHARIA, 2002. Floricultură generală, Ed. AcademicPres,
Cluj- Napoca.
7. CANTOR, MARIA, IOANA POP, 2005. Floricultură specială. Bază de date, Ed.
AcademicPres, Cluj- Napoca.
8. CIOCÂRLAN, V., M. BERCA, C. CHIRILĂ, I. COSTE, GH. POPESCU, 2004. Flora
Sagetală a României, Ed. Ceres, Bucureşti.
9. CIOCÂRLAN, V., 2008. Rectifying a few errors in the Romanian Exsiccate, J. Plant
Develop, 15, 27-28.
10. CIOCÂRLAN, V., 2009. Flora Ilustrată a României. Pteridophyta et Spermatophyta,
Editura Ceres, Bucureşti, pag. 833- 847.
Ghizela D. ALEC (VONICA) Summary of PhD thesis
22
11. COICIU, EVDOCHIA, G. RÁCZ, 1962. Plante medicinale şi aromatice, Ed. Academiei
R.P.R., Bucureşti.
12. DIHORU, GH., C. PÂRVU, 1987. Plante endemice în Flora României, Ed. Ceres,
Bucureşti.
13. DIHORU, GH., G. NEGREAN, 2009. Cartea Roşie a Plantelor Vasculare din România,
Ed. Academiei Române, Bucureşti.
14. DOSTÁL, J., 1976. Centaurea phrygia subsp. ratezatensis (Prod) Dostál, În: Greuter
W. (eds.), Euro + Med Plantbase, (2006 - 2009),
URL:[http://ww2.bgbm.org/EuroPlusMed, accessed 12 februar 2013].
15. DRĂGHIA, LUCIA, ALIONA MORARIU, LILIANA CHELARIU, 2010. Cytogenetic
research regarding species with ornamental value identified in the north-east area of
Romania, Cercetări Agronomice în Moldova, XLIII: 4, 144.
16. DRĂGULESCU, C-TIN, 1995. Botanica populară în Ţara Făgăraşului, Ed. Constant,
Sibiu.
17. DRĂGULESCU, C-TIN, 2008. Plante comestibile din România, Ed. Alma Mater,
Sibiu.
18. FLANN, CHRISTINA, W. GREUTER, D. J. HIND, 2010. Cassini’s Compositae genera:
A nomenclatural and taxonomic assesment, Taxon, 59, 1206- 1244.
19. FONT, MONICA G., 2007. Poiploidïa, filogénia I biogeografia en Centaurea L.
secció Acrocentron (Cass.) DC, Ph.D. Thesis, Facultat de Farmacia, Universitat de
Barcelona, Spain.
20. GARCIA-JACAS, NÚRIA, A. SUSANNA, 1992. Karyological notes on Centaurea sect.
Acrocentron (Asteraceae), Pl. Syst. Evol., 179, 1-18
21. GARCIA-JACAS, NÚRIA, A. SUSANNA, TERESA GARNATJE, ROSER VILATERSANA,
2001. Generic delimitation and phylogeny of the subtribe Centaurinae
(Asteraceae): a combined nuclear and chloroplast DNA analysis, Annals of Botany.,
87, 503-515.
22. GREUTER, W., 2006–2009. Compositae (pro major parte). – In: Greuter W. & Von
Raab-Straube E. (eds), Compositae. Euro+Med Plantbase – the information
resource for Euro - Mediterranean plant diversity,
URL:[http://ww2.bgbm.org/EuroPlusMed, accessed 12 februar 2013].
Ghizela D. ALEC (VONICA) Summary of PhD thesis
23
23. HAMMER, Ǿ., 1999-2012. Reference manual of PAST-Paleontological Statistics,
version 2.15, Natural History Museum and University of Oslo.
24. HAYEK, A., 1901. Die Centaurea - Arten Osterreich - Ungarns. Denkschr, Akad.
Wiss., 585- 783.
25. KOUTEKÝ, P., J. ŠTĔPÁNEK, TEREZA BADUROVA, 2012. Differentiation between
diploid and tetraploid C. phrygia: mating barriers, morphology and geographic
distribution, Preslia 84, 1-32.
26. LEPŠ, J., P. ŠMILAUER, 2003. Multivariate analysis of ecological data using
CANOCO, Cambridge Univ. Press, Cambridge.
27. NYÁRÁDY, E. I, 1943(a). Adatok a hazai Centaureák „Jacea” és „Phrygia”
csoportjainak bővebb ismeretéhez, különös tekintettel Kolzsvár környékének
flórájára. Additamenta ad cognitionem Centaurearum Hungariae, Senderabdruck
aus den „Botanikai Kozlemenyek”, Budapesta, XL: 1-2, 1-45
28. NYÁRÁDY, E. I, 1943(b). Űber unsere kritischen Centaureen mit berücksichtigung
von Wagners artikel, Borbasia, Tipografia Attila, Budapesta, 1-10.
29. PÉTIT, D., J. MATHEZ, A. QAID, 2001. Phylogeny of the Cardueae (Asteraceae)
based on analysis of morphological and palynological characters, Bocconea, 13, 41-
53.
30. PÎRVU, LUCIA, ALICE ARMATU, S. SCHIOPU, ILEANA RAU, D. COPREAN, 2008.
Centaurea cyanus L. - herba, chemical composition and therapeutical potential,
Proceeding of the International Symposium “New Research in Biotechnology”,
USAMV Bucharest, Romania, 187-194.
31. PRODAN, I., 1930. Centaureele României. Monografie, Instit. de arte grafice
„Ardealul”, Cluj- Napoca.
32. PRODAN, I., E. I. NYÁRÁDY, 1964. Centaurea L. – In: Săvulescu T. (eds.), Flora
Republicii Socialiste Romania, Edit. Academia R. S. Romania, Bucureşti. 9, 758–
951.
33. SÂRBU, ANCA, A. OPREA, I. SÂRBU, 2007. Plants from the habitat directive – Annex
IIb, presents in Romania, Buletinul Grădinii Botanice Iaşi, 14, 23-27.
Ghizela D. ALEC (VONICA) Summary of PhD thesis
24
34. SUSANNA, A., NURIA GARCIA-JACAS, D. E. SOLTIS, P. S.SOLTIS, 1995. Phylogenetic
relationships in tribe Cardueae (Asteraceae) based on ITS sequences, American
Journalof Botany, 82, 1056–1068.
35. SUSANNA, A., NÚRIA GARCIA-JACAS, 2007. Tribe Carduae, 123-146, În: Kadereit,
J.W. &C. Jeffrey, (Eds.), The Families and Genera of Vascular Plants, vol.8,
Flowering Plants. Eudicots. Asterales, Springer, Berlin & Heidelberg.
36. TEER BRAAK, C. J. F., P. ŠMILAUER, 2002. CANOCO reference manual and
CanoDraw for Windows user’s guide: software for canonical community ordination
(version 4.5). – Microcomputer Power, Ithaca, NY.
37. TUTIN, T. G., V.H. HEYWOOD, N. A. BURGES, D. M. MOORE, D. H. VALENTINE, S.
M. WALTERS, D. A. WEBB, 1964-1980. Flora Europaea, 1-5, Cambridge University
Press, Cambridge.
38. VONICA, GHIZELA, MARIA CANTOR, 2011 (a). The Polimorphism and Hybridization
of the Centaurea Species, Bulletin UASVM Horticulture, Cluj- Napoca, 68: 1, 444-
450.
39. VONICA, GHIZELA, MARIA CANTOR, 2011 (b). Some issues of Centaurea species
determination, Acta Musei Brukenthal, Sibiu, VI.: 3, 61-68.
40. VONICA, GHIZELA, MARIA CANTOR, 2012. Centaurea species attractive
Transylvania (Romania) endemic and rare plants with high potential as flowering
gardening plant, Bulletin UASVM Horticulture, Cluj-Napoca, 69: 1-2, 354-362.
41. VONICA, GHIZELA, ANDREEA TODORUŢ, A. S. BĂDĂRĂU, 2012. Morphological
variation of Centaurea atropurpurea Waldst. et Kit. species (subgenus Lopholoma)
from Transylvania, Acta Oecologica Carpatica, Sibiu, V, 41-56.
42. VONICA, GHIZELA D., P. KOUTECKÝ, MARIA CANTOR, B. I. HURDU, S. A.
BĂDĂRĂU, 2013 (a). New consideration regarding the endemic Centaurea
ratezatensis Prod from Romanian Carpathians, Symposium- Biogeography of the
Carpathians: Evolution of Biodiversity in a Spatiotemporal Context, Kraków
(Poland), 26th
- 28th
September 2013 (accepted paper).
43. VONICA, GHIZELA D., J. P. FRINK, MARIA CANTOR, 2013 (b). Taxonomic revision
of some taxa of Jacea - Lepteranthus group (Centaurea genus) based on
morphometric analysis, Brukenthal Acta Musei, Sibiu, VIII, in press.
Ghizela D. ALEC (VONICA) Summary of PhD thesis
25
44. WAGENITZ, G., F. HELLWIG, 1996. Evolution of characters and phylogeny of the
Centaurinae, 491- 510, În: D. J. N. Hind &Beentje (Eds.) Compositae: Systematics.
vol 1, Proceeding of the International Compositae Conference, Kew.
45. WAGNER, J., 1910. A magyarorszagi Centaureak ismertetése (Centaureae
Hungarieae), Math. és Terni. Közl., Budapesta, XXX: 6, 271-183.
46. ***,Flora Europaea- (http://rbg-web2.rbge.org.uk, 16 september 2011). Data
extracted from the digital version of the Flora Europaea. Royal Botanic Garden
Edinburgh, United Kingdom.
47. ***,Euro+Med, 2006, Euro+MedPlantBase - the information resource for Euro-
Mediterranean plant diversity. Available http://ww2.bgbm.org/EuroPlusMed/ [16
September 2011].
48. ***, HSIB- Ierbarul Muzeului de Istorie Naturală Sibiu- Muzeul Naţional
Brukenthal Sibiu.
49. ***, HCL- Ierbarul Gradinii Botanice “Alexandru Borza”- Univ. Babes-Bolyai
Cluj-Napoca.