8Blackwell Science, LtdOxford, UKBOJBotanical Journal of the Linnean Society0024-4074The Linnean Society of London, 2004? 2004
1453
K. ROMASCHENKO
345352
ET AL
Original Article
.
KARYOLOGY OF
CENTAUREA
Botanical Journal of the Linnean Society, 2004, 145, 345–352. With 24 figures
New chromosome counts in the Centaurea Jacea group
(Asteraceae, Cardueae) and some related taxa
KONSTANTYN ROMASCHENKO1, KUDDISI ERTUǦRUL, ALFONSO SUSANNA3*,
NÚRIA GARCIA-JACAS3, TUNA UYSAL2 and EMINE ARSLAN2
1
Institute of Botany M. G. Kholodny, Tereshchenkovska 2, 01106 Kiev, Ukraine
Department of Biology, Faculty of Science and Art, Selcuk University, Konya, Turkey
3
Botanic Institute of Barcelona (CSIC-Ajuntament de Barcelona), Passeig del Migdia s.n., E-08038
Barcelona, Spain
2
Received October 2003; accepted for publication February 2004
Twenty-seven chromosome counts are reported in 23 species of the genus Centaurea, mostly eastern endemic species
of the Jacea group, which has become the core of the re-defined genus Centaurea. Twenty reports are new, one is a
correction of a previous count, one is a confirmation of limited previous data and one represents a new basic number
in the Centaureinae. The prevalence of the basic chromosome number x = 9 among the Eastern sections of the Jacea
group is confirmed, together with the close correlation between karyological data and classification of the genus. Two
alternative hypotheses on the aberrant chromosome number (for the Centaureinae) found in C. behen are proposed.
One of them, if verified, would confirm that a cycle of polyploidy and descending dysploidy is a key mechanism in the
speciation of the group. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004,
145, 345–352.
ADDITIONAL KEYWORDS: Centaureinae – Compositae – Crimea – dysploidy – endemic – karyology –
Turkey.
INTRODUCTION
The taxonomy of the genus Centaurea L., with over
400 species, has always been rather complicated.
Recent surveys on the basis of DNA sequences and
morphological data (Garcia-Jacas et al., 2001; Susanna & Garcia-Jacas, in press) have circumscribed the
genus to species from sections Acrocentron DC. and
Cyanus (Mill.) DC., and the large assemblage of species of the Jacea group, which is the main subject of
this contribution.
The name ‘Jacea group’ indicates those taxa with
Jacea pollen type (Garcia-Jacas et al., 2000). This
group is the largest in the genus and shows an enormous diversity of habit, morphological adaptations,
morphology of the bract appendages, and cypselas.
There are three main characters that define the group.
Two are shared with sections Acrocentron and Cyanus: the lateral hilum on the seed (a derived character
*Corresponding author. E-mail: [email protected]
according to Dittrich, 1968) and the usual presence of
showy sterile peripheral florets lacking staminodes
(Wagenitz & Hellwig, 1996). The third character,
exclusive to the Jacea group, is the peculiar Centaurea
jacea pollen type, which differs from the rest of pollen
types by the very small size of the prolate or subprolate grains, and a caveate, microechinate or scabrid
exine (Wagenitz, 1955; Martín Villodre & GarciaJacas, 1999; Vilatersana et al., 2001).
Most of the taxa with this pollen type occur in the
eastern Mediterranean and the Irano-Turanian
regions. In previous reports on Turkish, Armenian and
Iranian species, we included species of the Jacea group
(Garcia-Jacas et al., 1997, 1998b; Garcia-Jacas, Susanna & Mozaffarian, 1998a). Our results suggested
that chromosome number is a good character in the
group, with a dysploid series ranging from x = 12 to
x = 7.
Having demonstrated the systematic potential of
karyology in the classification of Centaurea, together
with morphology and DNA sequence characteristics,
© 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145, 345–352
345
346
K. ROMASCHENKO ET AL.
our aims in this paper are to contribute to the general
knowledge of chromosome numbers in Centaurea,
especially in the Jacea group, of which few counts have
yet been made in eastern species, and to confirm the
relationships between systematics and karyology in
the sectional classification of the genus.
According to our data, this is the first chromosome
count for this species. This is a new confirmation of
x = 9 as the basic chromosome number of sect.
Acrolophus.
CENTAUREA
MATERIAL AND METHODS
Chromosome counts were made on somatic
metaphases using the squash technique. Root meristems from germinating seeds collected in the wild
were used.
Samples were pretreated with 0.002 M 8-hydroxyquinoline at 4∞C for 8 h. The material was fixed with
Carnoy for 24 h at low temperatures. Before staining,
the material was hydrolysed with 5N HCl for 1 h at
room temperature, stained with 1% acetic orcein and
mounted in 45% acetic acid. For all the counts, at least
five metaphase plates were examined from different
individuals. Preparations were made permanent by
freezing with CO2, dehydrating in ethanol and mounting in Canada balsam. Digital photographs were
taken using an Olympus 3030 camera mounted on an
Olympus microscope U-TV1 X. The preparations and
the herbarium vouchers are preserved in the Botanical Institute of Barcelona (BC) and the Department of
Biology, Faculty of Science and Art of the University of
Selcuk.
RESULTS AND DISCUSSION
CENTAUREA
SECT.
ACROCENTRON (CASS.) DC
Centaurea chrysantha Wagenitz
Turkey, Niǧde: Aladaǧlar, track above Demirkazık village, 2000 m, Ertu grul, Garcia-Jacas, Susanna 2298
& Uysal, 3.viii.2002 (BC). 2n = 18 (Fig. 1).
According to our data, this is the first chromosome
count for this species, a very narrow Turkish endemic.
As demonstrated in an earlier paper (Garcia-Jacas &
Susanna, 1992), x = 9 does not exist in Acrocentron.
The record of x = 9 in the Ibero-North African Centaurea clementei Boiss. by Humphries et al. (1978) was
wrong and the only known numbers in the section are
x = 10 and 11. This makes the position of C. chrysantha within Acrocentron very doubtful. Our molecular
data (A. Susanna, pers. observ.) also suggest that it
should be transferred to the Jacea group.
CENTAUREA
SECT.
ACROLOPHUS (CASS.) DC
Centaurea calolepis Boiss.
Turkey, Burdur-Muǧla: Dirmil mountain pass,
1600 m, Ertu grul, Garcia-Jacas, Susanna 2254 &
Uysal, 29.vii.2002 (BC). 2n = 18 (Fig. 2).
SECT.
CALCITRAPA DC
Centaurea alexandrina Delile
Egypt, Alexandria: 20 km west of Alexandria on the
coast road, Susanna 1854 & Vilatersana, 8.vi.1998
(BC). 2n = 20 (Fig. 3).
Our count disagrees with the 2n = 18 reported by
Kamel (1996). Most of the counts in this section suggest x = 10, as in the closely related sect. Tetramorphaea (DC). Boiss. (Garcia-Jacas et al., 1998a). There
are other incongruous counts in sect. Calcitrapa, however: x = 11 in C. hyalolepis Boiss. (Ghaffari & Chariat-Panahi, 1985; Ghaffari, 1989), together with
counts of 2n = 20 in the same species (Plitmann, 1976;
Bakhshi Khaniki, 1995), and 2n = 22 in C. pungens
Pomel (Reese, 1957; Hellwig et al., 1994). Other sections have at least two different basic chromosome
numbers (sects. Acrocentron, Cyanus and Mesocentron) and this could be the case in sect. Calcitrapa if
the position of C. pungens in the section is confirmed.
CENTAUREA
SECT.
CENTAUREA
Centaurea iconiensis Hub.-Mor.
Turkey, Konya: Seydisehir-Bozkır Yolu 20 km,
1050 m, Ertu grul 2480, 14.vii.2001 (herbarium
KNYA). 2n = 30 (Fig. 4).
This is the first count for this Turkish endemic. The
basic chromosome number x = 15 is the same as that of
most of the species of the section, in which there is
only one disparate record, of 2n = 26 in Centaurea centaurium L. (Bianco, D’Emerico & Medagli, 1990).
CENTAUREA
SECT.
CHEIROLEPIS (BOISS.) O. HOFFM.
Centaurea deflexa Wagenitz
Turkey, Konya: between Çukuryurt pass and Gevne
valley, 25 km from Taş kent, 1700 m, Ertu grul,
Garcia-Jacas, Susanna 2274 & Uysal, 1.viii.2002
(BC). 2n = 6x = 54 (Fig. 5).
As far as we know, this is the first count for this species. It is a hexaploid with x = 9. This high level is
widespread amongst the C. kotschyi complex as
C. kotschyi Hayek var. persica Wagenitz and
C. drabifolia Sm. ssp. detonsa (Bornm.) Wagenitz also
have 2n = 6x = 54 (Bakhshi Khaniki, 1995; GarciaJacas et al., 1997).
© 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145, 345–352
KARYOLOGY OF CENTAUREA
1
2
3
4
5
6
7
8
9
10
11
12
347
Figures 1–12. Somatic metaphases of Centaurea spp. Fig. 1. Centaurea chrysantha (2n = 18). Fig. 2. C. calolepis (2n = 18).
Fig. 3. C. alexandrina (2n = 20). Fig. 4. C. iconiensis (2n = 30). Fig. 5. C. deflexa (2n = 54). Fig. 6. C. kotschyi var. decumbens
(2n = 36). Fig. 7. C. pinardii (2n = 16). Fig. 8. C. aladaghensis (2n = 18). Fig. 9. C. cataonica (2n = 18). Fig. 10. C. kurdica
(2n = 18). Fig. 11. C. behen S-2321 (2n = 34). Fig. 12. C. behen S-2340 (2n = 34). In Figs 11 and 12 arrows indicate pairs of
larger chromosomes, each chromosome derived possibly from the fusion of two smaller ones. Scale bars = 10 mm.
© 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145, 345–352
348
K. ROMASCHENKO ET AL.
Centaurea kotschyi (Boiss. & Heldr.) Hayek var.
decumbens Wagenitz
Turkey, Konya: Fasikan Pass near Taş kent, 1800 m,
Ertu grul, Garcia-Jacas, Susanna 2279 & Uysal,
1.viii.2002 (BC). 2n = 4x = 36 (Fig. 6).
Centaurea kurdica Reichardt
Turkey, Elâzıǧ: Elâzıǧ to Bingöl road, 1 km from the
crossroad to Alacakaya, 700 m, Ertu grul, GarciaJacas, Susanna 2360 & Uysal, 6.viii.2002 (BC).
2n = 18 (Fig. 10).
This is the first count for this Turkish endemic variety.
It is a tetraploid, which makes the C. kotschyi group a
very interesting polyploid complex, as other varieties
of C. kotschyi are hexaploid (Bakhshi Khaniki, 1995).
Section Cheirolepis is indeed a polyploid group with
x = 9. No diploid record has been reported so far. Curiously, according to molecular data (Garcia-Jacas et al.,
2000), sect. Cheirolepis is connected to sect. Plumosipappus, whose only species, C. paphlagonica (Bornm.)
Wagenitz, has the same basic number and is diploid
with 2n = 18 (Garcia-Jacas et al., 1997).
According to our data, this is the first chromosome
count for this species.
With these three new counts in sect. Cynaroides, we
confirm that its basic chromosome number is x = 9.
The closely related sect. Paraphysis (DC). Wagenitz
also has the same number. There are two more diploid
counts of 2n = 18, C. amanicola Hub.-Mor. from
Turkey (Gardou & Tchehrehgocha, 1975) and
C. imperialis Haussk. ex Bornm. from Iran (GarciaJacas et al., 1998a; Ghaffari & Shahraki, 2001), and a
tetraploid one in C. charrellii Halácsy & Dörfler, an
endemic serpentinicole species of Greece (Constantinidis, Bareka & Kamari, 2002).
CENTAUREA
SECT.
CYANUS (MILL.) DC
Centaurea pinardii Boiss.
Turkey, Burdur: 4 km from Burdur on the road to
Isparta, outskirts of Askeriye, 950 m, Ertu grul,
Garcia-Jacas, Susanna 2244 & Uysal, 28.vii.2002
(BC). 2n = 16 (Fig. 7).
According to our data, this is the first chromosome
count for this species. Annual species of sect. Cyanus
have three different basic numbers, x = 8, 9 and 12
(Wagenitz & Hellwig, 1996). As suggested by GarciaJacas et al. (1997), variation in basic number in some
sections of Centaurea with dominance of annuals is
related to differences in their reproductive behaviour.
Autogamous species usually have higher, conserved
chromosome numbers than their allogamous relatives,
because autogamy precludes genetic interchange,
which seems to be essential for the occurrence of
descending dysploidy.
CENTAUREA
SECT.
CYNAROIDES BOISS.
EX
WALP.
Centaurea aladaghensis Wagenitz
Turkey, Adana: Aladaǧ supra Daǧdibi, 2000 m, Ertu grul, Garcia-Jacas, Susanna 2304 & Uysal, 3.viii.2002
(BC). 2n = 18 (Fig. 8).
According to our data, this is the first chromosome
count for this species.
Centaurea cataonica Boiss. & Hausskn.
Turkey, Gaziantep: Gaziantep, at the entry of the
town, 800 m, Ertu grul, Garcia-Jacas, Susanna 2319
& Uysal, 4.viii.2002 (BC). 2n = 18 (Fig. 9).
As far as we know, this is the first report for this
species.
CENTAUREA
SECT.
MICROLOPHUS (CASS.) DC
Centaurea behen L.
Armenia, Vaik: between Djermuk and the crossroad to
Goris, Fajvush, Gabrielyan, Garcia-Jacas, Guara,
Hovannisyan, Susanna 1562, Tamanian & Vallès,
21.viii.1995 (BC). 2n = 34.
Armenia, Abovian: between Gehart and Garni, Fajvush, Gabrielyan, Garcia-Jacas, Guara, Hovannisyan,
Susanna 1568, Tamanian & Vallès, 23.viii.1995 (BC).
2n = 36?,34.
Iran, Azarbayjan-e-Gharbi: 50 km west of Orumiyeh,
Garcia-Jacas, Mozaffarian, Susanna 1691 & Vallès,
6.viii.1996 (BC). 2n = 34.
Turkey, Maraş : 40 km from Gaziantep on the Mara ş
road, 2 km from Narlı, 500 m, Ertu grul, Garcia-Jacas,
Susanna 2321 & Uysal, 4.viii.2002 (BC). 2n = 34
(Fig. 11).
Turkey, Adiyaman: 5 km from Gölbaş i on the way to
Besni, 700 m, Ertu grul, Garcia-Jacas, Susanna 2340
& Uysal, 5.viii.2002 (BC). 2n = 34 (Fig. 12).
There are many counts for this species, all of them differing from ours (Chuksanova, Sveshnikova & Alexandrova, 1968; Tonjan, 1968; Avetisian & Tonjan,
1975; Ghaffari, 1986, 1989; Ghaffari & Shahraki,
2001). With the only exception of the doubtful record
by Chuksanova et al. (1968), who reported 2n = 26, all
the others were of n = 18 or 2n = 36. Ghaffari & Shahraki (2001) corrected their previous counts of n = 18–
2n = 32 + 0–2B, differing from previous results in
C. behen but more similar to other counts in the
section.
© 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145, 345–352
KARYOLOGY OF CENTAUREA
Our result shatters the karyological knowledge of
the section and indeed of the whole subtribe Centaureinae. This is the reason for our reticence in publishing it before gathering more evidence with the study of
more populations. Only in one of our Armenian collections did we find plates thought to be 2n = 36, but they
were unreliable prophases. When we repeated this
count, we found excellent metaphase plates with
2n = 34 and none with 2n = 36. Figures 11 and 12 demonstrate that there are no b-chromosomes that could
have accounted for this number, as suggested by Ghaffari & Shahraki (2001). All the chromosomes are normal, with centromeric constrictions.
Centaurea rigida Banks & Sol.
Turkey, Erzurum: 62 km from Erzurum on the road to
Bingöl, 10 km from Çat, Ertu grul, Garcia-Jacas,
Susanna 2377 & Uysal, 6.viii.2002 (BC). 2n = 16
(Fig. 13).
According to our data, this is the first chromosome
count for this species. Unlike that for C. behen, this
count agrees with those for many other species in
the section. Even though x = 8 is not as frequent as
x = 9, it still appears in some sections of the Jacea
group.
The unexpected result in C. behen confirms sect.
Microlophus to be aberrant within the Jacea group, in
which the dysploid series was previously limited from
x = 12 to 7. Indeed, the number x = 17 is aberrant
within the whole Centaureinae, where dysploidy was
limited from x = 16 to x = 7 (Garcia-Jacas, Susanna &
Ilarslan, 1996). However, x = 17 is a well-known basic
number in subtribe Carduinae (e.g. in Cirsium,
Cynara or Silybum; cf. Susanna & Garcia-Jacas, in
press). For this reason, we may conclude that C. behen
is in the process of reducing its chromosome complement from tetraploid (2n = 4x = 36) to neo-diploid
(2n = 2x = 34). In Figures 11 and 12 we have marked
with arrows a pair of chromosomes that are markedly
larger than the rest which could be the result of the
union of two smaller ones. The section would thus
have three basic numbers, x = 8, 9 and 17, because
there is a count of 2n = 18 in C. thracica (Janka)
Hayek (Damboldt & Matthäs, 1975; Constantinidis
et al., 2002).
An alternative explanation for the occurrence of
2n = 34 in C. behen is that it arose from the doubling of the chromosome number of an old hybridization between species with x = 8 and x = 9, both
numbers existing in sect. Microlophus. Centaurea
behen would be then a very widespread and fertile
allotetraploid. This hypothesis would also explain
the presence of the two larger chromosomes marked
with arrows in Figures 11 and 12, which would come
from the parent with the reduced chromosome num-
349
ber x = 8. Maybe the main difficulty for choosing
either of these possible origins is the problem of
tracking a hybridization that must be very old, given
the widespread distribution of the purported
allotetraploid.
CENTAUREA
SECT.
PHALOLEPIS (CASS.) DC
Centaurea sarandinakiae N. B. Illar.
Ukraine, Crimea: Planerskoe, Kara-Dag mountain,
Romaschenko, 16.viii.2002 (BC). 2n = 4x = 36 (Fig. 14).
According to our data, this is the first chromosome
count for this species.
Centaurea semijusta Juz.
Ukraine, Crimea: Simferopol, Chatyr-Dag mountain,
low plateau, Romaschenko, 1.ix.2002 (BC). 2n = 4x
= 36 (Fig. 15).
According to our information, this is the first chromosome count for this species.
Centaurea sterilis Stev.
Ukraine, Crimea: Planerskoe, Kara-Dag mountain,
Romaschenko, 16.viii.2002 (BC). 2n = 18 (Fig. 16).
According to our data, this is the first chromosome
count for this species.
Centaurea vankovii Klokov
Ukraine, Crimea: Alupka, Ai-Petri mountain, 1200 m,
Romaschenko, 30.viii.2002 (BC). 2n = 4x = 36 (Fig. 17).
According to our data, this is the first chromosome
count for this species.
Our counts confirm x = 9 to be the basic chromosome
number of sect. Phalolepis. All the studied species are
endemic in Crimea and most of them are known from
only a single locality. Our results also point out that
polyploidy has played a major role in the microspeciation of the section.
CENTAUREA
SECT.
PSEUDOPHAEOPAPPUS WAGENITZ
Centaurea antitauri Hayek
Turkey, Adana: Aladaǧ, above Daǧdibi, 2000 m,
Ertu grul, Garcia-Jacas, Susanna 2306 & Uysal,
3.viii.2002 (BC). 2n = 16 (Fig. 18).
Our result agrees with a report by Gardou & Tchehrehgocha (1975). This is a monotypic section without close relations within the Jacea group. The
relatively infrequent basic number x = 8 confirms
this isolation.
© 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145, 345–352
350
K. ROMASCHENKO ET AL.
13
14
15
16
17
18
19
20
21
22
23
24
Figures 13–24. Somatic metaphases of Centaurea spp. Fig. 13. Centaurea rigida (2n = 16). Fig. 14. C. sarandinakiae
(2n = 36). Fig. 15. C. semijusta (2n = 36). Fig. 16. C. sterilis (2n = 18). Fig. 17. C. vankovii (2n = 36). Fig. 18. C. antitauri
(2n = 16). Fig. 19. C. donetzica (2n = 36). Fig. 20. C. proto-gerberi (2n = 18). Fig. 21. C. pseudoleucolepis (2n = 18). Fig. 22.
C. cheirolepidioides (2n = 36). Fig. 23. C. cheirolopha (2n = 18). Fig. 24. C. isaurica (2n = 18). Scale bars = 10 mm.
© 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145, 345–352
KARYOLOGY OF CENTAUREA
CENTAUREA
SECT.
PSEUDOPHALOLEPIS KLOKOV
Centaurea donetzica Klokov
Ukraine, Donetzkaya: Krasny Liman, Romaschenko,
12.viii.2002 (BC). 2n = 4x = 36 (Fig. 19).
According to our data, this is the first chromosome
count for this species.
Centaurea proto-gerberi Klokov
Ukraine, Luganskaya: Stanichno-Lugansk,
aschenko, 5.ix.2002 (BC). 2n = 18 (Fig. 20).
Rom-
As far as we know, this is the first count for this
species.
Centaurea pseudoleucolepis Kleopow
Ukraine, Donetzkaya: Kamennye Mogily national reservation, Romaschenko, 1.viii.2002 (BC). 2n = 18
(Fig. 21).
According to our information, this is the first chromosome count for this species.
Our counts are the first for this section, that uniformly has x = 9.
CENTAUREA
SECT.
PSEUDOSERIDIA WAGENITZ
Centaurea cheirolepidioides Wagenitz
Turkey, Konya: Hadim, Gevne valley, 1500 m, Ertu grul 2268, 18.vii.2000 (herbarium KNYA). 2n = 4x = 36
(Fig. 22).
According to our data, this is the first chromosome
count for this species.
Centaurea cheirolopha (Fenzl) Wagenitz
Turkey, Maraş : 40 km from Gaziantep on the Mara ş
road, 2 km from Narlı, 500 m, Ertu grul, Garcia-Jacas,
Susanna 2324 & Uysal, 4.viii.2002 (BC). 2n = 18
(Fig. 23).
As far as we know, this is the first report for this
species.
Other counts in this section are 2n = 18 in C.
hermannii F. Herm. and C. lancifolia Spreng. (Hellwig, 1994) and 2n = 16 in C. stevenii M. Bieb. (Poddubnaja-Arnoldi, 1931). Our results support x = 9
as the basic number for sect. Pseudoseridia. Curiously, the contradictory report of 2n = 16 in
C. stevenii, if confirmed, would support a connection with sect. Rhizocalathium Tzev., which also
has x = 8 (Garcia-Jacas et al., 1998b), a connection
supported by ITS sequence analysis (Garcia-Jacas
et al., 2000).
INCERTAE
351
SEDIS
Centaurea isaurica Hub.-Mor.
Turkey, Karaman: Ayrancı, Avdan Daş ı, northern
stony slopes, 1500 m, Ertu grul 2311, 22.vii.2000 (herbarium KNYA). 2n = 18 (Fig. 24).
According to our data, this is the first chromosome
count for this Turkish endemic species. Sectional classification of this species is unclear (Wagenitz, 1975)
and it could be placed either in sect. Cheirolepis or
sect. Pseudoseridia.
CONCLUDING REMARKS
Our new counts confirm that x = 9 is the most frequent
chromosome number in the Eastern species of the
Jacea group, with five out of six sections of Eastern
distribution included in this paper having this number. If we extend these figures to the whole of the Eastern group of sections, 11 out of 18 have x = 9, three
have x = 8, two have x = 10 and one has x = 12. Only
one section, Ptosimopappus (Boiss.) O. Hoffm., is still
karyologically unknown.
As demonstrated repeatedly, correlation between
karyology and systematics is very close in Centaurea.
The case of C. chrysantha is paradigmatic: karyology
shows that it cannot be placed in Acrocentron and
instead suggests the Jacea group as its logical position, as confirmed on molecular grounds (A. Susanna
et al., unpubl. data).
Finally, if further studies confirm that the result of
x = 17 for C. behen is not the result of hybridization, it
could be the final proof of an old hypothesis. The cycle
of polyploidy and descending dysploidy is at the origin
of diversification in Centaurea (Garcia-Jacas et al.,
1996), and indeed is also the origin of the extreme dysploidy that is almost a trademark of the whole tribe
Cardueae (Susanna & Garcia-Jacas, in press).
ACKNOWLEDGEMENTS
This research was supported by the Dirección General
de Investigación, Ministerio de Ciencia y Tecnología
(project BOS2001-3041-C02-02), and the Generalitat
de Catalunya (‘Ajuts a Grups de Recerca Consolidats’
2001SGR00125). K. Romaschenko benefited from a
grant given by the NATO Programme in Spain. We
also thank Dr P. Brandham, from the Royal Botanic
Gardens, Kew, who critically read the manuscript and
made many suggestions for improving the text.
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