Botanical Journal of the Linnean Society, 2002, 139, 109–124. With 19 figures
Karyotaxonomy of Greek serpentine angiosperms
THEOPHANIS CONSTANTINIDIS1*, ELEFTHERIA-PERDIKO BAREKA2 and
GEORGIA KAMARI2
1
Institute of Systematic Botany, Department of Agricultural Biotechnology, Agricultural University of
Athens, Iera Odos 75, GR – 118 55 Athens, Greece
2
Section of Plant Biology, Department of Biology, University of Patras, GR – 265 00 Patras, Greece
Received October 2001; accepted for publication February 2002
The chromosome numbers are given for 20 angiosperm taxa, most of which grow predominately or exclusively on
serpentine in Greece, and the karyotype morphology is illustrated in all cases but one. Chromosome data are
provided for the first time for ten taxa (Alyssum pogonocarpum, Centaurea charrellii, C. vlachorum, Cephalaria
fanourii, Matricaria tempskyana, Onosma stridii, Scorzonera doriae, Silene fabarioides, S. salamandra, Trinia
glauca ssp. pindica) and for the endemic monospecific genus Leptoplax. A new number is reported for Thymus teucrioides ssp. candilicus. An unexpected dysploid number is recorded for a population of Leontodon hispidus ssp.
hispidus. Chromosome number and karyotype details from Greek populations are presented for the rest of the taxa.
Chromosomal evidence supports close relationships among members of Onosma subsect. Asterotricha. The unusual
chromosome number of Centaurea vlachorum supports its placement either in C. sect. Jacea or C. sect. Cyanus.
The evolution of taxa in sections Vierhapperia, Pulvinares, and Nervosae of Scorzonera appears to be connected with
particular chromosomal rearrangements and dysploidy. Leptoplax does not share the same chromosome number
with Peltaria but with Bornmuellera instead, something that facilitates infrageneric hybridization. Further issues
of taxonomy, distribution, evolution and conservation of serpentine species are briefly discussed where appropriate.
© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 139, 109–124.
ADDITIONAL KEYWORDS: chromosome numbers – distribution – endemism – Flora Hellenica – karyotype
morphology – ultramafic substrate.
INTRODUCTION
Rich in silica, magnesium and iron, the serpentine
rocks and soils (also sometimes called ophiolitic or
ultramafic) have long been known to support a specialized flora and to affect the form of vegetation
(Brooks, 1987; Baker, Proctor & Reeves, 1992; Roberts
& Proctor, 1992). Plant species growing on serpentine
may be morphologically distinct and in some cases
restricted to such geoedaphics, i.e. serpentine endemics. Endemic taxa whose taxonomically closely
related congener is easily recognized, often growing in
proximity to the serpentine endemic and occasionally
on a different rock type, are usually called neoendemics. A neoendemic may have undergone minimal
range expansion (Mayer & Soltis, 1994). Palaeoendemics can also been found on serpentine, especially
*Corresponding author. E-mail: [email protected]
when their relatives are not obvious or are distantly
distributed (Wild & Bradshaw, 1977; Brooks, 1987).
Palaeoendemics may be restricted to small areas as a
result of severe range contraction, leaving them often
with a disjunct distribution.
Certain cases of differentiation and evolution, occasionally with karyological data included, have been
investigated in serpentine endemics or species occupying both serpentine and nonserpentine substrates
(Westerbergh & Saura, 1992; Mayer, Soltis & Soltis,
1994; Dolan, 1995; Štepánková, 1996; Krahulcová &
Štepánková, 1998). Evidence derived from chromosome number and morphology may be useful in
tracing taxonomic and evolutionary relationships of
serpentine species, especially when information on
related species is also available. However, beyond
taxonomic description of certain taxa, very little work
has been devoted to the study of chromosome number,
karyotype morphology and possible evolutionary
© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 139, 109–124
109
110
T. CONSTANTINIDIS ET AL.
consequences of taxa with particular preference to
serpentine in Greece.
This paper aims to investigate chromosome number
and karyotype morphology of angiosperm taxa, most
of which grow predominately or exclusively on serpentine in Greece. It forms part of our studies on the
karyology of the Greek flora (Constantinidis, Kamari
& Phitos, 1997; Constantinidis & Kamari, 2000), and
on the various peculiarities of plant life on serpentine
(e.g. edaphic endemism and speciation, disjunct distributions, metal hyperaccumulation; Constantinidis,
1999; Psaras et al., 2000). The karyological information obtained is used in conjunction with published evidence on taxa related to those growing on serpentine
to address various taxonomic or evolutionary issues.
Eleven of our chromosome reports are published for
the first time, new chromosome numbers are given for
two taxa, karyotype morphology is given for the first
time in 13 taxa, and novel karyological records based
on Greek populations are provided for three taxa.
Details on the distribution and conservation of serpentine taxa have been added where appropriate.
cultivated in the Experimental Botanical Garden,
University of Patras. Taxa names, provenances and
chromosome numbers are summarized in Table 1.
The chromosome counts were made from root
tip metaphases. In all cases root tips were subject
to a chemical pretreatment (0.03% v/w aqueous 8hydroxyquinoline) and/or chilling (4°C) for 3–24 h
before being processed further. Details of the squash
technique used are reported in Constantinidis et al.
(1997). Chromosome terminology principally follows
Levan, Fredga & Sandberg (1964) and Stebbins (1971).
Nomenclature for the taxa examined mostly follows
the published volumes of Med-Checklist (Greuter,
Burdet & Long, 1984, 1986, 1989) or, in genera not
appearing in them or revised more recently, Tutin
et al. (1976, 1993), Strid (1986a), Strid & Tan (1991a)
and Greuter (1997). Investigated taxa are listed by
their families, in alphabetical order.
RESULTS AND DISCUSSION
BORAGINACEAE
Onosma elegantissima Rech. fil. & Goulimy
MATERIAL AND METHODS
Living plant material of perennial taxa and seeds of
annual species was collected in various serpentine
areas of Greece. Individual plants (3–8 per taxon) were
Nomos Grevenon, Eparchia Grevenon: Lower slopes
of Mt. Vourinos, c. 2.5–3.3 km after Dafneron village along road to Exarchos. Serpentine slopes with
Juniperus oxycedrus and Buxus sempervirens. Alt.
Table 1. Alphabetical list of the taxa investigated (E = endemic to Greece) with provenance and chromosome number
found: = first report; = first record based on Greek material; = new chromosome number; + = first report of
karyotype morphology
Taxon
Provenance
2n
Alyssum pogonocarpum
Centaurea charrellii (E)
Centaurea thracica
Centaurea vlachorum (E)
Cephalaria fanourii (E)
Crepis merxmuelleri (E)
Hesperis laciniata ssp. laciniata
Leontodon hispidus ssp. hispidus
Leptoplax emarginata (E)
Matricaria tempskyana (E)
Onosma elegantissina (E)
Onosma stridii (E)
Ranunculus pedatus
Scorzonera doriae
Silene fabaria ssp. domokina (E)
Silene fabarioides
Silene haussknechtii (E)
Silene salamandra (E)
Thymus teucrioides ssp. candilicus (E)
Trinia glauca ssp. pindica
Rodos island
Domokos area, Sterea Ellas
Mt. Katachloron area, Sterea Ellas
Mts. Aftia/Flega, Ipiros
Mt. Gramos, Ipiros/Makedonia
Mt. Smolikas, Ipiros
Mt. Smolikas, Ipiros
Mt. Smolikas, Ipiros
Domokos area, Sterea Ellas
Mt. Zigos, Ipiros
Mt. Vourinos, Makedonia
Mt. Kallidromon, Sterea Ellas
Domokos & Mt. Kallidromon, Sterea Ellas
Eptachori area, Makedonia
Domokos area & Mt. Gerania, Sterea Ellas
Mt. Mavrovouni, Ipiros
Mt. Smolikas, Ipiros
Rodos island
Domokos area, Sterea Ellas
Mt. Aftia/Flega, Ipiros
16 36 18 24 +
18 +
8+
12 12 16 +
36 +
14 +
14 +
16 12 +
24 +
24 +
24 +
24 +
28 18 +
© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 139, 109–124
KARYOTAXONOMY OF GREEK SERPENTINE ANGIOSPERMS
c. 720–750 m, Lat. 40°10¢ N, Long. 21°36¢ E, 30.iv.2000,
Constantinidis 9010 (UPA). 2n = 14 (Fig. 1).
Known only from a single mountain in north central Greece (Vourinos) and growing from c. 700 up to
1800 m, this attractive species has been found in
several localities forming multiple rosettes on rather
shallow, serpentine ground. We have not observed it
growing on limestone but Teppner (1991a) gave it as
such. This being true, Onosma elegantissima should
not be considered as an obligate serpentine endemic.
In agreement with Strid (1995) and Strid & Tan (2000)
we considered it as hardly threatened. Most populations encountered appeared to be healthy, untouched
by browsing animals (except perhaps for some missing
flowering shoots), and with a good number of juvenile
individuals.
A report of the O. elegantissima chromosome
number is given by Teppner (1991a, 1991b) as 2n = 14,
without further details. Our investigation confirmed
the same diploid number. The chromosomes are
mostly metacentric. Satellites were observed on two
of the smallest pairs. The chromosome formula of
the population examined is given as 2n = 10 m +
2 m-SAT + 2 m/sm-SAT = 14. Examination of mitotic
plates at metaphase and especially at late prophase
revealed clear heterochromatic bands, mostly situated
at centromeric and distal parts of the longer chromosome arms. The number and size of chromosomes and
the rather late condensing euchromatin support placement of O. elegantissima complement with the O.
echioides L. cytotype, characteristic of asterotrichous
Onosma species with x = 7 (Teppner, 1971, 1991b).
When originally described (Rechinger, 1957),
Onosma elegantissima was compared to the Albanian
endemic O. mattirolii Bald. Judging from the published karyological information on the latter species
(Baltisberger & Baltisberger, 1995) some differences
from C. elegantissima are noted. In particular, clear
submetacentric chromosomes exist that we did not
find in O. elegantissima and do not match an O.
echioides cytotype.
Several karyotypes of members of O. subsect. Asterotricha (Boiss.) Gürke studied so far with 2n = 14
present a remarkable similarity, which is of probable taxonomic importance, especially if the diversity
of chromosome number and morphology observed
in Onosma is taken into account. Indeed, the karyotype of O. elegantissima, a single mountain endemic,
appears almost identical to those of O. echioides from
Italy (Teppner, 1971; Raimondo, Rossitto & Ottonello,
1983), the Greek endemic O. erecta Sm. ssp. erecta
(Teppner, 1988a), the single mountain endemic O.
stridii (see below), and O. inexspectata Teppner, an
endemic species of SW Anatolia (Teppner, 1974). This
stable and conservative karyotype feature may actually represent a symplesiomorphy shared by species
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distributed quite differently. Karyological information
on many Asiatic species is still lacking.
Onosma stridii Teppner
Nomos
Fthiotidos,
Eparchia
Lokridos:
Mt.
Kallidromon, c. 2.7 km after the village of Modion,
along road to Reginion. Slopes with scrub and a dry
streambed. Vegetation composed mainly of Cistus
salviifolius, Quercus coccifera, Pistacia lentiscus,
Juniperus oxycedrus, etc. Ophiolithic substrate. Alt. c.
580–620 m, Lat. 38°40¢ N, Long. 22°41¢ E, 22.iv.1998,
Constantinidis 7522 (UPA). 2n = 14 (Fig. 2).
Onosma stridii, a very local species known from a
small part of the Mt. Kallidromon serpentine area,
should be considered as an obligate serpentine stenoendemic. It was described relatively recently by
Teppner (1988b) from a single collection made in 1987.
Our search on Mt. Kallidromon located only two
populations: the one mentioned above (locus classicus) and a second to the north-north-west of it that
forms part of the understorey of locally sparse Pinus
halepensis woodland. This second population is composed of individuals with the usual white or whitish
flowers and others with pink, pinkish or tinged pink
flowers, a colour that is unique among Greek Onosma
species.
Karyological examination of the locus classicus
population revealed a diploid karyotype with 2n = 14.
The chromosomes are mostly metacentric, with at
least two very obvious satellites situated on the short
arm of the smallest chromosome pair. Another pair of
satellites is sometimes observed on a metacentric
chromosome pair. Therefore, the karyotype formula of
the population examined is given as 2n = 10 m +
2 m-SAT + 2 m/sm-SAT = 14. This is the first report on
the chromosome number and karyotype morphology of
O. stridii.
Together with O. elegantissima, O. stridii belongs
to the asterotrichous Onosma species characterized
by a base number of x = 7, a diploid cytotype and a
clear separation of euchromatin and heterochromatin at prophase or even early metaphase stages
(echioides type). The taxonomic affinity between the
two species that was assumed by Teppner (1988b) is
further supported by karyological evidence. Although
O. stridii has pollen larger than that of O. elegantissima, they are both diploid with very similar karyotype morphology.
CARYOPHYLLACEAE
Silene fabaria (L.) Sm. ssp. domokina Greuter
Nomos
Attikis/Viotias,
Eparchia
Megaridos/
Korinthias: northern foothills of Mt. Gerania, between
Pefkogiali and Mavrolimni settlements. Ophiolitic
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112
T. CONSTANTINIDIS ET AL.
1
2
3
4
5
6
7
8
Figures 1–8. Photomicrographs of mitotic metaphases. Scale bars = 10 mm. Fig. 1. Onosma elegantissima (2n = 14). Fig. 2.
O. stridii (2n = 14). Fig. 3. Silene fabaria ssp. domokina (2n = 24). Fig. 4. S. fabarioides (2n = 24). Fig. 5. S. haussknechtii
(2n = 24). Fig. 6. S. salamandra (2n = 24). Fig. 7. Centaurea thracica (2n = 18). Fig. 8. C. charrellii (2n = 36).
© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 139, 109–124
KARYOTAXONOMY OF GREEK SERPENTINE ANGIOSPERMS
substrate. Alt. 5–10 m, Lat. 38°03¢ N, Long. 23°07¢ E,
09.ix.1994, Constantinidis 3238 (UPA). 2n = 24
(Fig. 3).
Nomos Fthiotidos, Eparchia Domokou: c. 3.8 km
south of Domokos, along a secondary road to an abandoned quarry. Rocky slopes. Ophiolithic substrate.
Alt. c. 510 m. Lat. 39°05¢ N, Long. 22°18¢ E, 28.vi.1998,
Constantinidis (seeds collected). 2n = 24.
Described very recently by Greuter (1995), this
neoendemic taxon shows a disjunct distribution
between the serpentine parts around the town of
Domokos and the north-west serpentine slopes of Mt.
Gerania. Although it occurs predominately on serpentine, it was also observed growing on flysch, usually in
secondary, man-made habitats such as scree along
road embankments and disturbed alluvia.
Its chromosome number, 2n = 24, was reported by
Melzheimer (1987) under the name S. fabaria ssp.
thebana (Orph. ex Boiss.) Melzh. Of the two populations examined by Melzheimer only that from
Domokos is considered to represent the true ssp.
domokina, while the population from Thiva (Thive)
was later reclassified as ssp. fabaria (Greuter, 1995).
The same chromosome number of 2n = 24 is confirmed
here, based on material from two disjunct populations.
Chromosomes are mostly metacentric and submetacentric, with at least three pairs of clearly visible
satellites situated on the short arm of the longest
metacentric/submetacentric chromosome pair and on
the long arms of two metacentric pairs.
Based on the provided locality data, all previous
chromosome numbers for Silene fabaria s.l. that
appear in the literature (Damboldt & Phitos, 1968;
Degraeve, 1980; Baltisberger & Aeschimann, 1988)
should be attributed to S. fabaria ssp. fabaria. This
last taxon is distributed in east Greece, Aegean islands
and Turkey, mostly in coastal, rocky places and on a
variety of substrates (Melzheimer & Ulrich, 1994;
Greuter, 1997).
Silene fabarioides Hausskn.
Nomos
Ioanninon,
Eparchia
Metsovou:
Mt.
Mavrovouni, c. 15.5 km along forest road on the
mountain. The road starts c. 2.5 km after (north-west)
Metsovon (turnoff to Milea). Rocky slopes with Pinus
close to a stream. Mainly ophiolite. Alt. 1350–1400 m,
Lat. 39°51¢ N, Long. 21°05¢ E, 18.vi.2000, Constantinidis 9136 (UPA). 2n = 24 (Fig. 4).
A Balkan endemic species, Silene fabarioides is
often found on serpentine in Greece but is not
confined to it (Melzheimer, 1980; T. Constantinidis,
pers. observ.). Its distribution includes Albania, F.Y.R.
Macedonia, Bulgaria and Greece (Greuter, 1997).
Its chromosome number and morphology are presented here for the first time. The species is diploid
113
with 2n = 24, the most common number in Silene. Most
of the chromosomes are metacentric, with only a
few being metacentric/submetacentric. Three pairs of
satellites were observed, two on the short arms and
one on the long arms of metacentric chromosomes.
Silene haussknechtii Hausskn.
Nomos Ioanninon, Eparchia Konitsis: Mt. Smolikas,
c. 6.2 km along forest road starting just south-west
of the village of Pades and leading to the upper parts
of the mountain. Openings of Pinus forest and road
sides, serpentine. Alt. c. 1580 m, Lat. 40°03¢ N, Long.
20°54¢ E, 01.ix.1997, Phitos, Kamari & Constantinidis
25608 (UPA). 2n = 24 (Fig. 5).
Earlier literature records, specimens examined
and personal observations indicate that Silene
haussknechtii is a serpentine endemic. It has been
considered to be restricted to serpentine of north-west
Greece but its occurrence in serpentine areas of
adjacent Albania cannot be ruled out (Greuter, 1997).
Its chromosome number has been reported as
2n = probably 24, ranging between 22 and 25
(Oxelman, 1995) in material from the foothills of Mt.
Vasilitsa. Our count confirms the number of 2n = 24
and at the same time the karyotype morphology of
the species is presented. Most of the chromosomes
are metacentric, with two pairs of satellites on the
short arms of the longest submetacentric pair and a
medium-sized metacentric chromosome pair. In some
metaphase plates a spherical formation has been
found, almost equalling the short arm of some chromosome pairs in size. It might be a B-chromosome,
possibly partly responsible for the earlier difficulties
in estimating the exact chromosome number of the
species.
Silene salamandra Pamp.
Nomos Dodekanisou, Eparchia Rodou: Rodos island,
c. 4.4 km north-east of the village of Laerma. Sparse
Pinus brutia forest, scree and dry stream beds, ultramafic substrate. Alt. c. 320 m, Lat. 36°11¢ N, Long.
27°55¢ E, 18.iv.1998, Constantinidis K214 (UPA).
2n = 24 (Fig. 6).
A rare and local endemic of Rodos island, Silene
salamandra is an attractive annual species of S.
sect. Atocion Otth. Its taxonomic position has been
elucidated by Carlström (1986a), who simultaneously
recorded it as an exclusive endemic of the ophiolithic areas of Rodos (Carlström, 1986b). However,
the altitudinal range of the species (0–500 m) given by
Greuter (1997) indicates that it should also grow by
the sea, where the ophiolithic substrate does not exist.
Ultramafic rocks are found in the central parts
of Rodos island, presumably having chemical and/
or petrological synthesis differing from the similar-
© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 139, 109–124
114
T. CONSTANTINIDIS ET AL.
looking formations on the Greek mainland and Evvia.
They are rather sparsely covered with Pinus brutia
forest. During our visits on the island, S. salamandra
was observed growing on gravelly and rather steep
hill slopes, unstable scree, and dry streambeds with
coarse ultramafic sand. The species achieves a luxurious growth under cultivation in normal soil, and
increases its size up to six times compared to wild
specimens.
Chromosome number and karyotype morphology of
S. salamandra are reported here for the first time. All
chromosomes appear to be metacentric. Satellites are
not usually observable, with the exception of a distinct, spherical one on a chromosome of average size.
The same chromosome number of 2n = 24 has been
found in S. insularis Barbey (Damboldt & Phitos,
1966), a close relative of S. salamanda endemic to the
island of Karpathos. The diploid number of 24 is,
however, a very common and conservative feature in
Silene (although not in S. subg. Conoimorpha), and
uniformity of karyotype does not permit conclusions of
karyotaxonomic importance to be made.
COMPOSITAE
Centaurea charrellii Halácsy & Dörfler
Nomos Fthiotidos, Eparchia Domokou: Road LamiaDomokos, very close to the crossing towards Omvriaki
village. Margins of cultivated land and among bushes,
ophiolithic substrate. Alt. c. 500 m, Lat. 39°06¢ N,
Long. 22°18¢ E, 26.vi.1997, Constantinidis 6965 (UPA).
2n = 36 (Fig. 8).
One of the rarest endemic species of Centaurea in
Greece, C. charrellii is currently known from two very
small and severely threatened populations in the area
of Domokos. Both populations do not exceed 40 individuals in total and the species has justifiably been
characterized as Endangered (Georgiadis, 1995). Since
its original collection in the locus classicus close to
Vodena, now the city of Edessa (north Greece), it has
not been reported again from the same area, nor any
other area of Greece besides Domokos.
Owing to its tall habit, bright yellow florets (superficially resembling the much more common C. salonitana Vis. from a distance) and taxonomic/chorological
position, i.e. the only representative of the Asiatic C.
sect. Cynaroides Boiss. ex Walp. in Greece and Europe,
without any close relatives, and growing more than
1200 km from any other member of the same section
(Wagenitz, 1975a) the species itself is distinct and
unique. While collecting around Domokos we found C.
charrellii twice, growing on serpentine or on a mixture
or serpentine and flysch, usually under the protection
of grazed Quercus coccifera scrub. Contrary to previous reports (Georgiadis, 1995), we found that the
species is able of reproducing itself sexually. All the
capitula of the plants examined contained viable
achenes that germinated under cultivation, 2 months
after being collected, without presenting any signs of
dormancy. As, however, breeding systems and reproductive ecology of C. charrellii are virtually unknown,
there is always the possibility that seed setting is prevented in very small populations, especially in cases
in which the species is self-incompatible. Normal
sexual reproduction, viable seeds, progeny that are
identical to their parents, and lack of related species
in Greece make the assumption of Georgiadis (1995)
that C. charrellii might be of hybrid origin improbable, unless it is an ancient allotetraploid derived from
the doubling of the chromosome number of a hybrid
between two diploid species.
The chromosome number and a microphotograph of
a metaphase plate of C. charrellii are presented here
for the first time. The species is tetraploid, with
2n = 36. The somewhat solid appearance of the chromosomes does not permit a detailed morphological
description, but at least metacentric and submetacentric chromosome types occur in the complement. Four
satellites were also observed.
The chromosome number of 2n = 36 is clearly based
on x = 9, a common basic number in Centaurea found
in several sections (e.g. Hellwig, 1994; Garcia-Jacas,
Susanna & Mozaffarian, 1998a; Garcia-Jacas et al.,
1998b). As far as C. sect. Cynaroides is concerned, two
other members have been investigated karyologically,
C. amanicola Hub.-Mor. from Turkey (Gardou &
Tchehrehgocha, 1975) and C. imperialis Boiss. ex
Walp. from Iran (Garcia-Jacas et al., 1998a). They are
both diploid with 2n = 18. As examination of relevant
literature did not reveal chromosome numbers in any
other species of the more than 22 members of C. sect.
Cynaroides, ours is the first report of tetraploidy in the
section.
Centaurea thracica (Janka) Hayek
Nomos Karditsis, Eparchia Karditsis: Mt. Katachloron,
along road from Kedros village to Loutra Smokovou,
c. 2.6 km south of Kedros. Meadow, rocky slopes and
road sides. Ophiolithic substrate. Alt. c. 220 m, Lat.
39°11¢ N, Long. 22°02¢ E, 27.vi.1997, Constantinidis
7061 (UPA). 2n = 18 (Fig. 7).
By analogy to Centaurea charrellii, C. thracica is the
only representative of the Asiatic sect. Microlophus
(Cass.) DC. in Europe, but it is not as rare as C. charrellii. We have always collected or observed C. thracica
on serpentine in the area of Domokos, the foothills of
Mt. Katachloron (the population examined karyologically), the medium-altitude parts of Mt. Kratsovon
and between the villages of Kakoplevri and Oxinia,
north-west of Kalambaka. A record from Evvia island
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KARYOTAXONOMY OF GREEK SERPENTINE ANGIOSPERMS
(Rechinger, 1961) also strengthens this correlation.
However, exclusive conjunction of this species with
ultramafic substrates still has to be proved.
The diploid chromosome number found, 2n = 18, is
a confirmation of an earlier report by Damboldt &
Matthäs (1975), on material obtained from the botanical garden of Sofia, Bulgaria. This is the first time
that the chromosome number of a Greek population is
examined. Most of the chromosomes are metacentric
or submetacentric, with two satellites visible on the
short arm of a submetacentric chromosome pair and
two on the shorter arms of a metacentric chromosome
pair. This is generally in close agreement with the
karyotype drawing provided by Damboldt & Matthäs
(1975).
As already noted by Garcia-Jacas et al. (1998b)
C. sect. Microlophus is characterized by two basic
numbers, x = 8 and x = 9. It is not clear yet to what
extent those different numbers are related to the taxonomy and phylogeny of its species, as to our knowledge, chromosome numbers of only three out of more
than ten species comprising the section are known
currently.
Centaurea vlachorum Hartvig
Nomos Ioanninon, Eparchia Dodonis/Metsovou: Mts.
Aftia/Flega, c. 16.5 km after turn to Milea of the road
Ioannina-Trikala and along a secondary road that
leads to the upper parts of the mountain. Mixed
Fagus-Pinus forest and small clearings. Ophiolite. Alt.
c. 1650 m, Lat. 39°52¢ N, Long. 21°06¢ E, 28.vii.1999,
Constantinidis 8669 (UPA). 2n = 24 (Fig. 9).
Centaurea vlachorum is a rare endemic species
known from only two mountains, Milea and Aftia of
Northern Pindos (Hartvig, 1981). It has always been
found associated with serpentine and can therefore be
considered as a serpentine endemic. Since its description, the systematic placement of the species has been
intriguing. C. vlachorum is considered to be a distinct
species without close relatives and an isolated member
of C. sect. Jacea (Mill.) DC. (in its widely ascribed
sense, including sect. Lepteranthus DC.; see Wagenitz,
1975b, 1980), although morphological features of
bracts are reminiscent of sect. Cyanus (Mill.) DC. and
Acrocentron (Cass.) DC. (Gamal-Eldin & Wagenitz,
1991). The chromosome number of the species, previously unknown, would elucidate its sectional placement, given that several groups of Centaurea have
quite characteristic base numbers.
C. vlachorum is diploid with 2n = 24. Chromosomes
are of average size and mostly metacentric. Satellites
are found on the shorter arms of a metacentric and
a submetacentric pair. It is characteristic that one
pair of chromosomes often has a condensation rate
differing from that of the rest of the complement,
115
evident by its banding effect in metaphase plates.
Differentiation of staining may reflect heterochromatin rich areas on this particular submetacentric
chromosome pair. The karyotype formula of the population examined is given as 2n = 18 m + 2 mSAT + 2sm + 2 sm-SAT.
The diploid karyotype of C. vlachorum is based on
x = 12, a number intermediate between the more primitive x > 12 and the more advanced x < 12 groups of
Centaureinae (Garcia-Jacas, Susanna & Ilarslan,
1996). Within Centaurea, x = 12 is confirmed in three
sections, namely Mesocentron (Cass.) DC. (species of
sect. Hymenocentron (Cass.) DC. are often included
here), Cyanus, Jacea and Hyalaea DC. Based on the
available chromosome evidence, no relationship
between C. vlachorum and C. sect. Acrocentron is
supported. All members of sect. Acrocentron so far
examined karyologically have the chromosome base
of x = 10 or x = 11 (Garcia Jacas & Susanna de la
Serna, 1992; Routsi & Georgiadis, 1999), with x = 12
essentially unknown in the section. Furthermore,
morphological characters clearly exclude any close
relationship between C. vlachorum and members of
sections Mesocentron/Hymenocentron and of the small
section Hyalaea. In sect. Cyanus, x = 12 is known in
C. cyanus L. and C. montana L. (several counts in
Fedorov, 1969) while in sect. Jacea it is found in, e.g.
C. phrygia L. ssp. nigriceps (Dobrocz.) Dostál (see
Fedorov, 1969), in which confirmation of the count
would be welcome.
To conclude, karyological evidence indicates that the
isolated placement of C. vlachorum in C. sect. Jacea
(contradicted by bract morphology) is in accordance
with its unusual chromosome number. The same
number exists in sect. Cyanus, but the placement of
C. vlachorum in this section is not in agreement with
its pollen morphology.
Crepis merxmuelleri Kamari & Hartvig
Nomos Ioanninon, Eparchia Metsovou: Mt. Smolikas,
southern slopes of the mountain, c. 5.7 km along
road from Pades village to Armata village. Rocky
places by the road, scree and road sides. Ophiolitic
substrate. Alt. c. 1050 m, Lat. 40°02¢ N, Long. 20°56¢ E,
03.ix.1997, Constantinidis 8634 (UPA) and Phitos,
Kamari & Constantinidis 26819 (UPA). 2n = 8
(Fig. 10).
Crepis merxmuelleri is a rare species, described
recently from the serpentine belt of Mt. Smolikas
(Kamari & Hartvig, 1988). The description was based
on two specimens collected on the north-east parts of
the mountain, at an altitude of c. 1250–1550 m. Our
collections, found on the south-east part of the same
mountain and at a slightly lower altitude, extend
somewhat the known distribution of the species.
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116
T. CONSTANTINIDIS ET AL.
9
10
11
12
13
Figures 9–13. Photomicrographs of mitotic metaphases. Scale bars = 10 mm. Fig. 9. Centaurea vlachorum (2n = 24). Fig. 10.
Crepis merxmuelleri (2n = 8). Fig. 11. Leontodon hispidus ssp. hispidus (2n = 12). Fig. 12. Scorzonera doriae (2n = 12);
arrowheads indicate secondary constrictions of the longest chromosome pair. Fig. 13. Matricaria tempskyana (2n = 36).
Compared to the isotype (Hartvig & Seberg 5104,
UPA!), the original description and a recent collection
(Constantinidis & Kouki 9876, ACA!, UPA!), our specimens present a significant difference: they completely
lack the yellowish-brown glandular indumentum,
which has been attributed to C. merxmuelleri as a key
character (Kamari, 1991).
The chromosome number of 2n = 8 has already been
reported for C. merxmuelleri (Kamari, 1991). The
number is confirmed here, and a detailed karyotype
analysis is given for the first time. The species is
diploid with submetacentric, acrocentric, and subtelocentric chromosomes. Small satellites are observed on
the small arm of the subtelocentric chromosome pair.
© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 139, 109–124
KARYOTAXONOMY OF GREEK SERPENTINE ANGIOSPERMS
The karyotype formula is given as 2n = 2 sm + 4 st +
2 t-SAT = 8.
Overall morphology, chromosome number, and karyotype analysis of C. merxmuelleri are strongly reminiscent of C. turcica Degen & Bald. (see Constantinidis
& Kamari, 2000; for a presentation of the karyotype
of C. turcica and for references). The two species
resemble each other in that they both have glandular
and eglandular forms in different populations. It
seems that they are closely related species differing in
their substrate preferences. C. turcica grows on limestone and rarely on flysch, while C. merxmuelleri has
been found so far only on serpentine.
Leontodon hispidus L. ssp. hispidus
Nomos Ioanninon, Eparchia Konitsis: Mt. Smolikas,
close to the small lake known as Drakolimni, along
road from a pen situated c. 10 km north of Pades
village. Stony slopes, scree and rocky places. Ophiolitic
substrate. Alt. c. 2000–2200 m. Lat. 40°05¢ N, Long.
20°54¢ E, 22.vi.1998, Phitos, Kamari & Constantinidis
cult. 9137 (UPA). 2n = 12 (Fig. 11).
Although Leontodon hispidus ssp. hispidus is a
widespread taxon having broad ecological preferences
with respect to the geological substrate, the karyotype
examination of a population growing on the serpentine
parts of Mt. Smolikas is included here because of its
unexpected chromosome number. All five plants examined and 13 microphotographs of metaphase plates
taken confirm the chromosome number of 2n = 12.
This differs from the many previous counts of 2n = 14
for L. hispidus, reported from different countries, e.g.
Bulgaria (Kuzmanov & Georgieva, 1976), the British
Isles (Morton, 1977), Finland (Uotila & Pellinen,
1985), Spain (Izuzquiza & Nieto Feliner, 1991) and
Greece (Strid & Franzén, 1981). There are also some
rare reports of triploid plants and some aneuploid
cases with 2n = 16 or 18 (De Groot, 1977) but, to our
knowledge, the number 2n = 12 is new for this taxon.
Given the unusual chromosome number counted, the
specimens were again checked morphologically but no
significant deviation from other forms of L. hispidus
was found, bearing in mind the variability observed in
this taxon.
The karyotype formula of the Smolikas population
is found as 2n = 2 m + 6 sm + 2 sm-SAT + 2 st-SAT = 12.
In some plates one of the metacentric chromosomes,
the smallest in size in the complement, tends to be
submetacentric. The acrocentric satellited chromosome pair is the longest in the complement, while the
submetacentric one is the shortest. Our formula is
close to the general classification of chromosomes
reported by Rousi (1973) and Pittoni (1974), but the
incidence of acrocentric (st) chromosomes given by
117
Rousi (1973) is greater (two–three pairs), compared
with our results (one pair).
The dysploid reduction of chromosome number from
2n = 14 to 2n = 12 seems to be related to the loss of a
chromosome pair, nevertheless additional chromosomal translocations cannot be ruled out. Relative size
of chromosomes in the serpentine population, when
compared with published information from different
countries, does not support chromosome fusion. As
Arabidopsis-type sequence repeats (TTTAGGG)n have
been traced only at telomeric positions in L. hispidus
with 2n = 14 (Fuchs, Brandes & Schubert, 1995), it
would be of interest to check the 2n = 12 population using fluorescent in situ hybridization (FISH) for
similar signals, in an effort to trace possible chromosome rearrangements. It is still unknown whether or
not such a chromosome change was induced by the
serpentine soil in which L. hispidus grows.
Matricaria tempskyana (Freyn. & Sint.) Rauschert
[Syn. Tripleurospermum tempskyanum (Freyn &
Sint.) Hayek]
Nomos Ioanninon, Eparchia Metsovou: Mt. Zigos,
c. 1.2 km after ‘Rachi Kataras’ along road to the
telecommunication station. Wooded slopes and roadsides. Ophiolithic substrate. Alt. c. 1780 m, Lat.
39°46¢ N, Long. 21°13¢ E, 07.vii.1999, Constantinidis
8634 (UPA). 2n = 36 (Fig. 13).
This Greek endemic species is found in montane
areas of north-west Greece, where it grows mostly
on serpentine (Strid & Tan, 1991b). In two populations examined morphologically, the one cited above
and another growing in Pinus forest margins close to
the village of Milea, the vast majority of individuals
were lacking outer ligulate florets but in some cases
these were present in a much-reduced, rudimentary
form.
The tetraploid chromosome number of 2n = 36 and
the karyotype morphology for Matricaria tempskyana
are given here for the first time. Three chromosome
types were observed in the complement: metacentric
and submetacentric chromosomes are dominating,
while acrocentric ones are limited to eight pairs.
Four small satellites mark the short arm of a submetacentric and an acrocentric chromosome pair. The
karyotype formula of the population examined is given
as 2n = 14 m + 2 m/sm + 10 sm + 2 sm-SAT + 6 st +
2 st-SAT = 36.
Scorzonera doriae Degen & Bald.
Nomos Kastorias, Eparchia Kastorias: c. 5.0–5.8 km
south of Eptachori village. Serpentine slopes, rocks
and scree by the road. Alt. c. 840 m, Lat. 40°13¢ N,
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T. CONSTANTINIDIS ET AL.
Long. 20°58¢ E, 09.vii.1999, Constantinidis 8683
(UPA). 2n = 12 (Fig. 12).
Scorzonera doriae is endemic to the western Balkan
Peninsula. In Greece, it is known from a few localities,
all on serpentine of the North Pindos area (see Lack
& Kilian, 1991 for mountain reports).
The chromosome number and karyotype morphology
of S. doriae are presented here for the first time. The
species is diploid with 2n = 12. Five out of six chromosome pairs in the complement are metacentric,
while the longest sixth pair is submetacentric, characterized by a satellite on the short arm. The karyotype
formula is given as 2n = 10 m + 2 sm-SAT = 12.
The longest chromosome pair of S. doriae is of
particular interest. A wide, poorly stained centromeric
area separates the long from the short arm in each
chromosome, and one or two secondary constrictions
may be visible on each long arm (Fig. 12). A closer
examination of other members in S. sect. Vierhapperia Lipsch., to which S. doriae belongs, reveals that S.
hirsuta L. from Spain (Díaz de la Guardia & Blanca,
1987) and probably S. biebersteinii Lipsch. from the
Caucasus area (Nazarova, 1997) have a karyotype
morphology similar to that of S. doriae. Long, submetacentric chromosome pairs are not usually found
in Scorzonera members with 2n = 14, where chromosomes bearing satellites are among the smallest in
the complement. A comparison of the chromosome
idiograms given by Nazarova (1997) is even more
informative. Long, submetacentric chromosome pairs
bearing a satellite are also found in n = 6 karyotypes
of sections Pulvinares (Boiss.) Lipsch. and Nervosae
Lipsch. Remarkable similarities in chromosome structure between S. hirsuta and S. hispanica L. have
already urged Díaz de la Guardia & Blanca (1987) to
propose that each long submetacentric chromosome in
S. hirsuta may be the result of a reciprocal translocation combining two smaller, metacentric chromosomes. Here, the same procedure of metacentric
(usually marked with satellites) chromosome fusion in
Scorzonera members with n = 7 to yield long, submetacentric chromosomes in Scorzonera members
with n = 6 is given a more general perspective. If n = 7
in Scorzonera is accepted as ancestral (Nazarova,
1997), then n = 6 is derived and secondary. Chromosomal rearrangements such as chromosome fusion
should have played an important role in the evolution
of particular Scorzonera sections. Obviously, they complement the general evolutionary scheme proposed by
Nazarova (1997).
CRUCIFERAE
Alyssum pogonocarpum A. Carlström
Nomos Dodekanisou, Eparchia Rodou: Rodos island,
c. 4.4 km north-east of the village of Laerma. Sparse
Pinus brutia forest, scree and dry stream beds, ultramafic substrate. Alt. c. 320 m, Lat. 36°11¢ N, Long.
27°55¢ E, 18.iv.1998, Constantinidis K 219 (UPA).
2n = 16.
Endemic to Rodos island and adjacent Turkish
mainland (Carlström, 1987), Alyssum pogonocarpum
was found growing on open ultramafic scree and
gravel of central Rodos, together with other interesting species such as Aethionema arabicum (L.) O. E.
Schulz, Cleome iberica DC., Linum virgultorum
Planchon, and Silene salamandra. Its chromosome
number of 2n = 16 is reported here for the first time.
The somewhat blurred appearance of chromosomes
in metaphase plates, however, did not permit their
illustration.
Alyssum pogonocarpum was compared with A.
hirsutum, considered to be its closest ally (Carlström,
1984). The two species, in addition to morphological divergence, also exhibit different chromosome
numbers. Alyssum pogonocarpum is diploid with
2n = 16, while A. hirsutum is polyploid with 2n = 46
or 48 (see Ančev & Dudley, 1981, and reference in
Goldblatt & Johnson, 1994).
Hesperis laciniata All. ssp. laciniata
Nomos Ioanninon, Eparchia Konitsis: Mt. Smolikas,
the south-east lower parts of the mountain, c. 3.4 km
after the village of Distraton along road to Samarina.
Ophiolithic scree. Alt. c. 1050 m. Lat. 40°02¢ N, Long.
21°00¢ E, 11.viii.1998, Constantinidis & Garofalo
(seeds collected, voucher: K242, UPA). 2n = 12
(Fig. 14).
Most populations of Hesperis laciniata are found
on calcareous soils, usually in rocky places, gravel and
scree. Populations growing on serpentine occur more
rarely, at least in Greece. Morphological comparison of
serpentine and limestone collections of H. laciniata
ssp. laciniata did not reveal any noteworthy
differences.
The chromosome number of 2n = 12, found in
three plants of H. laciniata ssp. laciniata from Mt.
Smolikas, is the result of the first karyological
examination of a Greek population. The karyotype
analysis revealed the formula of 2n = 6 m +
2 m-SAT + 2 sm + 2 sm/st-SAT = 12, with satellites
usually difficult to observe on the contracted chromosomes of metaphase plates. This diploid number is
in accordance with the reports given by Ančev &
Goranova (1997) for H. laciniata ssp. laciniata and by
Constantinidis & Kamari (1994) for H. laciniata ssp.
secundiflora (Boiss. & Spruner) Breistr. Small differences between the above-cited reports and this study
centre on the relative position of the satellites and the
classification of one chromosome pair as submetacentric or subtelocentric. The gametic number of n = 6,
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KARYOTAXONOMY OF GREEK SERPENTINE ANGIOSPERMS
14
15
16
17
18
19
119
Figures 14–19. Photomicrographs of mitotic metaphases. Scale bars = 10 mm. Fig. 14. Hesperis laciniata ssp. laciniata
(2n = 12). Fig. 15. Leptoplax emarginata (2n = 16). Fig. 16. Cephalaria fanourii (2n = 18). Fig. 17. Thymus teucrioides ssp.
candilicus (2n = 28). Figure 18. Ranunculus pedatus (2n = 16). Fig. 19. Trinia glauca ssp. pindica (2n = 18).
given by Ruíz de Clavijo (1994) for material from
Spain should be better attributed to H. l. ssp.
spectabilis (Jord.) Rouy & Foucaud and not to H. l. ssp.
laciniata, especially if the work on the genus by
Dvořák, 1971) is taken into account.
Leptoplax emarginata (Boiss.) O. E. Schulz
Nomos Fthiotidos, Eparchia Domokou: c. 1 km south
of Ekkara village along road to Ano Agoriani village.
Stony places by a small stream. Ophiolithic substrate.
Alt. c. 200 m, Lat. 39°08¢ N, Long. 22°11¢ E, 26.vi.1997,
Constantinidis 7020 (UPA). 2n = 16 (Fig. 15).
Leptoplax O. E. Schulz is usually considered an
independent, monotypic genus (e.g. Hartvig, 1986a),
although its sole species may occasionally be found in
the bibliography as belonging to Peltaria Jacq., under
the name of P. emarginata (Boiss.) Hausskn. Leptoplax
is endemic to Greece, distributed from the area of
North Pindos to the north, down to Mt. Gerania in the
south. The genus is also found close to the border with
Albania, where is has not yet been reported (see
Paparisto et al., 1988), so it should be sought there.
Leptoplax is always associated with serpentine and
can be considered as a serpentine endemic.
The chromosome number of 2n = 16 and a microphotograph of the karyotype of Leptoplax are reported
here for the first time. The species is diploid. Chromosomes are small and appear to be either metacentric or submetacentric but clear identification is
© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 139, 109–124
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T. CONSTANTINIDIS ET AL.
difficult. A satellite appears on one metacentric chromosome but its homologue is not usually visible.
Leptoplax does not have the same chromosome
number as Peltaria, in which several counts have
given the stable number of 2n = 14 or the tetraploid
2n = 28 (see Fedorov, 1969; Van Loon, 1987 for
references). This difference in base chromosome
number supports the splitting of Leptoplax from
Peltaria, although there are cases in Cruciferae
where more than one chromosome base-number
exists in the same genus. In contrast, Leptoplax has
the same chromosome number as Bornmuellera
Hausskn., something that may facilitate hybridization, pollen fertility in hybrids and introgression
between species of the two genera, a process that
has already been observed in nature and reported by
Hartvig (1986a).
DIPSACACEAE
Cephalaria fanourii Perdetzoglou & Kit Tan
Nomos Ioanninon/Kastorias, Eparchia Konitsis/
Kastorias: Mt. Gramos, upper parts of Bouchetsi
summit, c. 100–400 m north-north-west of the highest
summit. Steep, east-facing, treeless serpentine slopes
and scree. Alt. c. 1680 m, Lat. 40°14¢ N, Long. 20°57¢ E,
13.ix.2000, Constantinidis & Kyparissis 9150 (UPA).
2n = 18 (Fig. 16).
A rare serpentine endemic, Cephalaria fanourii is
known, up to now, from a single locality, the serpentine top area of Mt. Bouchetsi, where it has been collected independently by P. Hartvig and colleagues, and
by D. Perdetzoglou (Perdetzoglou & Tan, 1995).
According to Perdetzoglou & Tan (1995), C. fanourii
belongs to C. sect. Leucocephalae. This section is a
member of C. subgen. Denticarpus, which supposedly
includes species whose achenes have eight apical
teeth (‘achenium apice dentibus 8 minoribus vel longioribus’, see Szabó, 1940). This feature obviously does
not apply to C. fanourii or to its relatives (as defined
by Perdetzoglou & Tan, 1995) C. laevigata (Waldst. &
Kit.) Schrader and C. coriacea (Willd.) Steud. In C.
fanourii there are four small, apical teeth on the
achene, alternating with four minute ones that may
even be absent. C. laevigata achenes have no apical
teeth at all, while those of C. coriacea also lack teeth
or may rarely have minute ones. Thus, the character
of apical teeth on the achene allows reliable distinction between C. fanourii and its related taxa.
The chromosome number of C. fanourii, reported
here for the first time, is 2n = 18, the most common
number in Cephalaria (Verlaque, 1985). Karyotype
morphology is given as 2n = 8 m + 2 m-SAT + 2 sm +
2 sm-SAT + 4 st = 18. The satellites are large and
clearly visible. This chromosome formula does not
match any of the Cephalaria karyotype formulae given
by Verlaque (1985) and adds to the karyotypic heterogeneity of the genus.
LABIATAE
Thymus teucrioides Boiss. & Spruner ssp. candilicus
(Beauv.) Hartvig
Nomos Fthiotidos, Eparchia Domokou: c. 5 km south
of Ekkara village along road to Ano Agoriani village.
Stony slopes, field margins and road sides. Ophiolithic
substrate. Alt. c. 460 m, Lat. 39°07¢ N, Long. 22°12¢ E,
26.vi.1997, Constantinidis 7005 (UPA). 2n = 28
(Fig. 17).
This is the second chromosome count of Thymus
teucrioides ssp. candilicus, a Greek neoendemic taxon
apparently found exclusively on serpentine. The first
count, originating from a population growing on the
serpentine parts of Mt. Gerania, gave the number of
2n = 30 (Constantinidis et al., 1997). The new count of
2n = 28 adds a different chromosome number to this
taxon and simultaneously addresses the case of dysploidy in Thymus. Unfortunately, karyotype morphology is not possible to determine from the Domokos
population. Lack of chromosome details is due mostly
to the small size of chromosomes in Thymus, and does
not allow any direct karyotype comparison among
populations.
To our knowledge, no other chromosome counts
are known in Th. sect. Teucrioides besides those of
Thymus teucrioides ssp. candilicus. Different chromosome numbers in the same Thymus species, similar
to the case reported here, are not exceptional. For
example, Th. praecox Opiz has been found to have
2n = 54, 56–58 (Jalas & Pohjo, 1965 as T. humifusus
Bernh.; Trela-Sawicka, 1972) and Th. glabrescens
Willd. (= Th. odoratissimus Miller) has 2n = 28, 32, 56,
58 (Trela-Sawicka, 1968). A cytogeographical investigation of Th. sect. Teucrioides in Greece, where it is
largely distributed, might be of interest.
RANUNCULACEAE
Ranunculus pedatus Waldst. & Kit.
Nomos Fthiotidos, Eparchia Domokou: About 5.2 km
south-south-east of Ekkara village, along road to
Ano Agoriani. Slopes with Quercus coccifera, Juniperus oxycedrus and Cistus spp., small meadows and
margins of fields. Ophiolithic substrate. Alt. c.
580–620 m, Lat. 39°07¢ N, Long. 22°12¢ E, 18.iii.1998,
Constantinidis 7386 (UPA). 2n = 16 (Fig. 18).
Nokos Fthiotidos, Eparchia Lokridos: Mt.
Kallidromon, c. 8.3 km after the village of Elatia, along
forest road to Reginion. Slopes with Quercus coccifera,
Arbutus andrachne, Pistacia lentiscus, etc. Ophiolithic
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KARYOTAXONOMY OF GREEK SERPENTINE ANGIOSPERMS
substrate. Alt. c. 630 m, Lat. 38°40¢ N, Long. 22°46¢ E,
23.iv.1998, Constantinidis 7595 (UPA).
A rare species in Greece, Ranunculus pedatus was
known with certainty until now from a single collection on the ophiolithic summit area of Mt. Kratsovon
in the North Pindos area (Strid, 1986b). Our collections from serpentine areas extend the distribution
of the species in Greece further south, and to a lower
altitude. It is expected that more populations will
be found if serpentine areas are carefully searched in
early spring.
The chromosome number of 2n = 16 is counted on
Greek material of R. pedatus for the first time. The
two populations examined present a similar karyotype, which has the following formula: 2n = 6 m +
2 sm + 2 sm-SAT + 2 st + 2 st-SAT + 2 t-SAT = 16. The
satellites are small. The chromosome number is in
agreement with reports from other countries, e.g.
Slovakia (Májovský et al., 1978). The tetraploid
number of 2n = 32 is also reported for this species (see
Goldblatt, 1988 for reference). The karyotype formula
of R. pedatus given here conforms to the general
karyotype scheme of R. sect. Ranunculastrum presented by Goepfert (1974).
UMBELLIFERAE
Trinia glauca (L.) Dumort ssp. pindica Hartvig
Nomos Ioanninon, Eparchia Dodonis/Metsovou: Mts.
Aftia/Flega. c. 16.5 km after turn to Milea of the road
Ioannina-Trikala and along a secondary road that
leads to the upper parts of the mountain. Mixed
Fagus-Pinus forest and small clearings. Ophiolite. Alt.
c. 1650 m, Lat. 39°52¢ N, Long. 21°06¢ E, 28.vii.1999,
Constantinidis 8663 (UPA). 2n = 18 (Fig. 19).
Originally described as a distinct taxon by Hartvig
(1986b) and distributed in Greece and Albania, Trinia
glauca ssp. pindica is known to grow both on serpentine and limestone substrates of several Greek
mountains. Its chromosome number and karyotype
morphology are presented here for the first time.
This taxon is diploid with 2n = 18. The karyotype
consists mostly of metacentric chromosomes, with a
pair of very clear satellites marking a metacentric/
submetacentric chromosome pair. The karyotype
formula can be described as 2n = 12 m + 2 m/sm +
2 m/sm-SAT + 2 sm = 18.
The chromosome number of 2n = 18 in Trinia glauca
ssp. pindica agrees with several chromosome reports
on T. glauca ssp. glauca from various European countries (Kieft & Van Loon, 1978; Váchová & Lhotská,
1978; Váchová & Lhotska, 1980; Van Loon, 1980;
Bellomaria & Hruška, 1983; Nikolov, 1991) or Trinia
glauca s.l. (Van Loon & Van Setten, 1982; Castroviejo
& Nieto Feliner, 1986). Karyotype illustration in
121
Trinia glauca is not usually given or, when given,
does not permit an unambiguous classification of
chromosomes.
CONCLUSIONS
In certain cases, karyological information did not
provide evidence of taxonomic importance between
serpentine species and their relatives, as, e.g. in sections of Silene, where chromosome number and karyotype morphology tend to be conservative features.
Differences were found in other groups of taxa, and
were referring to ploidy level (diploid Alyssum pogonocarpum vs. polyploid A. hirsutum), chromosome number (between the genera Leptoplax and Peltaria)
or karyotype morphology (Onosma elegantissima vs.
O. mattirolii). Differences also exist among populations of the same taxon (dysploidy in a serpentine
population of Leontodon hispidus ssp. hispidus). These
are indications of karyotype differentiation and may
reflect taxonomic relationships between serpentine
species and their allies that are more distant than
were considered previously. For Onosma, Crepis and
Leptoplax in particular, new taxonomic relationships
are proposed, which are in concordance with karyological data and observations in the field.
In other examples, chromosome number and
karyotype morphology of serpentine taxa are related
to evolutionary modifications characterizing supraspecific taxonomic units (e.g. common karyotype
morphology of Onosma subsect. Asterotricha, chromosomal rearrangements in Scorzonera sects. Vierhapperia, Pulvinares and Nervosae). These shared
evolutionary features may be irrelevant to the geological substrate on which plant taxa grow today, or may
precede adaptation and specialization on serpentine.
Our results so far indicate that karyotype changes
in serpentine taxa, compared to their close allies,
follow no strict modification patterns. The reaction of
plants to the serpentine factor may vary, and concentration on particular taxonomic groups in the future
will afford more precise information on karyotypic
differentiation enhanced by serpentine.
ACKNOWLEDGEMENTS
This work was partially supported by a grant from the
Greek Ministry for Development, General Secretariat
for Research and Technology (Project EPET II, no.
9513514).
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