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A karyosystematic study of the genus Bellevalia Lapeyr. (Hyacinthaceae) in Greece


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The genus Bellevalia is represented in Greece by eight taxa, three of which are endemic. Bellevalia brevipedicellata (2n = 8) and B. sitiaca (2n = 16) are restricted to the island of Kriti, while B. hyacinthoides (2n = 8, 12) is distributed in the Kiklades Islands, the central and southern mainland and the Ionian Islands. Four taxa, i.e. B. dubia subsp. boissieri (2n = 8), B. trifoliata (2n = 8), B. romana (2n = 8) and B. ciliata (2n = 8, 16) are Mediterranean elements. The presence of B. edirnensis (2n = 24) is reported as new for the Greek flora. New ploidy levels of three Bellevalia species (triplo-, tetra- and hexaploids) are reported for the first time. The main morphological features, the chromosome numbers, the karyotype morphology, as well as the geographical distribution and further issues of taxonomy and conservation of all Bellevalia taxa in Greece are presented and discussed. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739.
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A karyosystematic study of the genus Bellevalia Lapeyr.
(Hyacinthaceae) in Greece
Botanical Institute, Section of Plant Biology, Department of Biology, University of Patras, GR – 265
00 Patras, Hellas (Greece)
Received 2 January 2008; accepted for publication 7 February 2008
The genus Bellevalia is represented in Greece by eight taxa, three of which are endemic. Bellevalia brevipedicellata
(2n=8) and B. sitiaca (2n=16) are restricted to the island of Kriti, while B. hyacinthoides (2n=8, 12) is distributed
in the Kiklades Islands, the central and southern mainland and the Ionian Islands. Four taxa, i.e. B. dubia subsp.
boissieri (2n=8), B. trifoliata (2n=8), B. romana (2n=8) and B. ciliata (2n=8, 16) are Mediterranean elements.
The presence of B. edirnensis (2n=24) is reported as new for the Greek flora. New ploidy levels of three Bellevalia
species (triplo-, tetra- and hexaploids) are reported for the first time. The main morphological features, the
chromosome numbers, the karyotype morphology, as well as the geographical distribution and further issues of
taxonomy and conservation of all Bellevalia taxa in Greece are presented and discussed. © 2008 The Linnean
Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739.
ADDITIONAL KEYWORDS: chromosome numbers – conservation – distribution – endemism – karyology –
polyploidy – taxonomy.
Bellevalia Lapeyr. is an attractive genus of the
Hyacinthaceae family and consists of small perennial
geophytes interesting from both a taxonomic and
karyological point of view.
The genus comprises over 65 taxa, distributed in
the Mediterranean and the central-west Asiatic area,
while the largest number of species is located in the
Irano-Turanian region (Feinbrun, 1938–1940).
In flowering material, the most useful morphologi-
cal character for distinguishing the genus Bellevalia
from its closely related genera is the shape of the
filaments. These are fused throughout the whole
length of the perianth tube. The free parts are more
or less triangular, flat and usually connate at the base
(Persson & Wendelbo, 1979). Closely related genera,
such as Muscari Mill., Hyacinthella Schur. and Hya-
cinthus L., all have thread-like filaments.
The morphological differentiation within Bellevalia
taxa is rather weak. However, for the classification at
species level, the main characters traditionally used
are: leaf width and pubescence, raceme shape and
density, pedicel/perianth length ratio, tube/lobe ratio
and perianth colour, anther and bud features (Cowley,
Özhatay & Mathew, 1994), as well as seed morphology.
Until recently, seven taxa of the genus had been
recorded, in Greece, three of which are endemic, i.e.
Bellevalia hyacinthoides (Bertol.) K. Persson & Wen-
delbo, B. brevipedicellata Turrill and B. sitiaca Kyp-
riotakis & Tzanoudakis, while the remaining four, B.
dubia (Guss.) Kunth subsp. boissieri (Freyn) Feinbrun,
B. trifoliata (Ten.) Kunth, B. romana (L.) Sweet and B.
ciliata (Cyr.) Nees. are Mediterranean elements.
In 2004, some plants were collected at the edge of
cultivated fields in Evros province (north-eastern
Greece), which lie very close to the borders with
Edirne villayet of European Turkey. These are mor-
phologically different from any other Greek Bellevalia
taxa known so far. The examination of morphological
features and the determination of chromosome
number lead us to conclude that the plants from
*Corresponding author. E-mail:
Botanical Journal of the Linnean Society, 2008, 157, 723–739. With 20 figures
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739 723
north-eastern Greece belong to B. edirnensis N.
Özhatay & Mathew, one of the most localized endem-
ics of European Turkey (Özhatay et al., 1991b) and
now new for the flora of Greece.
Dried Bellevalia material was studied from ATH,
ATHU, E, ISTE, LINN, NAP, UPA and W Herbaria
(abbreviations according to Holmgren & Holmgren,
2007), as well as from the Museum of Natural History
of Kefalonia-Ithaki (in this paper abbreviated as
Living plants from 54 investigated populations
were cultivated in the experimental garden of the
Botanical Institute, University of Patras, for karyo-
logical studies. These populations are indicated by an
asterisk in the specimen list of the Appendix.
The chromosome counts were obtained from root-tip
metaphases, using the squash technique (Östergren
& Hennen, 1962). Root tips were pretreated for 6 h
in a mixture of 1 : 1 8-hydroxyquinoline (0.002%
w/v) : colchicine 0.2% w/v and followed by fixation in
Carnoy (3 : 1 (v/v) absolute ethanol : glacial acetic
acid) for 24 h at 0–4 °C. Afterwards, they were
hydrolysed in 1N HCl for 12 min at 60 °C and placed
in Feulgen’s stain (Darlington & La Cour, 1969) for
c. 2–4 h. At least five photomicrographs of each popu-
lation were examined, taken with a Zeiss Axiophot
photomicroscope. Chromosome terminology follows
Levan, Fredga & Sandberg (1964), Stebbins (1971)
and Kamari (1976), taking into consideration com-
ments and suggestions given by Sybenga (1959),
Benzer et al. (1971) and Favarger (1978).
1. Bellevalia dubia (Guss.) Kunth, Enum. Pl. 4: 308
(1843) Hyacinthus dubius Guss., Cat. H. Reg.
Boccad.: 32 & 78 (1821). – Lectotype (Borzatti de
Loewenstern & Garbari, 2003: 544) (Italy, Sicily)
‘Palermo’, April, Gussone (NAP, photo!).
According to Feinbrun (1938–1940) the species is
divided into three subspecies: subsp. dubia, subsp.
hackeli (Freyn) Feinbrun, and subsp. boissieri
(Freyn) Feinbrun. Of these subspecies, only B.
dubia subsp. boissieri occurs in Greece.
Bellevalia dubia subsp. boissieri (Freyn) Fein-
brun, Palest. Jour. Bot. (Jerusalem) 1(4): 348
(1940) Bellevalia boissieri Freyn, Flora 68: 95
(1885). – Syntypes (several) from Greece: ‘in insula
Hydra’, Heldreich, Pichler and from Dalmatia: ‘in
insula Lesina’, Jirusch (Herb.?) (Fig. 1).
Leaves 3–6, 18–60 cm ¥3–8.5 mm, equalling or longer
than scape, procumbent, lingulate, canaliculate,
margins smooth. Scapes 1–3, 13–37 cm. Raceme 12-
to 45-flowered, cylindrical. Pedicels (2.5–)3–6 mm,
ascending or horizontal. Flower buds blue. Perianth
4–7 mm, campanulate, bluish–violet at anthesis, with
white green-veined lobes. Anthers violet. Fruiting
raceme cylindrical, pedicels shorter than the capsule,
ascending or horizontal. Valves of capsule broadly
obovate, emarginate at apex. Seeds broadly ellipsoid
to globular. – 2n=8, 12 chromosomes.
Flowering: March to the end of May.
Habitat: It grows in moist banks, grassy places and
cultivated ground.
Distribution: Central and eastern parts Mediterra-
nean region: Croatia, Italy, Albania, Greece, Turkey.
In Greece it is distributed mainly in Sterea Hellas,
Peloponnisos and the Ionian Islands.
According to Feinbrun (1938–1940), the typical sub-
species of B. dubia is distributed in S Italy, including
Sicily, the subsp. hackeli occurs in Portugal and
1. Perianth tube shorter than the lobes; seeds pyriform....................................................8. B. hyacinthoides
Perianth tube equalling or longer than the lobes; seeds broadly ellipsoid to globular....................................2.
2. Raceme conical at anthesis..................................................................................................4. B. ciliata
Raceme cylindrical, ellipsoid to cylindrical or oblong at anthesis...............................................................3.
3. Anthers yellow.............................................................................................................5. B. edirnensis
Anthers violet..................................................................................................................................4.
4. Leaves procumbent; perianth bluish–violet with white green-veined lobes..............1. B. dubia subsp. boissieri
Leaves erect or recurved; perianth not bluish–violet with white green-veined lobes......................................5.
5. Pedicels longer than 3 mm.................................................................................................................6.
Pedicels shorter than 1.5 mm..............................................................................................................7.
6. Raceme cylindrical; perianth tubular–campanulate with tube violet at the base........................2. B. trifoliata
Raceme oblong; perianth turbinate with tube whitish at the base ............................................3. B. romana
7. Leaves undulate with obscure parallel veins; valves of capsule obovoid to elliptic...........6. B. brevipedicellata
Leaves not undulate, with visible parallel veins; valves of capsule sub-orbicular ..........................7. B. sitiaca
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
subsp. boissieri in the central and eastern Medi-
terranean region. Recently, Borzatti de Loewenstern
& Garbari (2002, 2003) consider that the typical sub-
species is an endemic unit of the Sicilian flora, while
the plants of south Italy belong to subsp. boissieri.
Besides the accepted combination B. dubia (Guss.)
Kunth, the variations B. dubia (Guss.) Rchb. and B.
dubia (Guss.) Roemer & Schultes are also frequently
used. However, these combinations should be treated
with suspicion and are probably incorrect, as Reichen-
Figures 1–8. Bellevalia taxa occurring in Greece. Fig. 1, B. dubia subsp. boissieri. Fig. 2, B. trifoliata. Fig. 3, B. romana.
Fig. 4, B. ciliata. Fig. 5, B. brevipedicellata. Fig. 6, B. sitiaca. Fig. 7, B. hyacinthoides. Fig. 8, B. edirnensis.
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
bach (1830), to whom Roemer & Schultes also refer,
used for the description of the species a specimen
collected ‘In Ligurien bei Pontremoli’, where only
B. webbiana Parl. has been recorded (Borzatti de
Loewenstern & Garbari, 2003).
2. Bellevalia trifoliata (Ten.) Kunth, Enum. Pl. 4:
308 (1843) Hyacinthus trifoliatus Ten., Fl. Nap.
3: 376, t. 136 (1824). Type: (Italy) ‘in arvis Apuliae,
Foggia’, Tenore (iso- FI, not seen) (Fig. 2).
=Hyacinthus abortivus Caval., Deux Nouv. Esp.:
14 (1848) B. abortiva (Caval.) Gren., Fl. Fr. 3:
217 (1855).
=B. pendulina Chiov., Bull. Soc. Bot. Ital.: 283
Leaves 3–5, (8–)10–42 cm ¥(5–)9–27 mm, equalling or
longer than scape, erect, lingulate, margins scabrid
or smooth. Scapes 1–3, 23–55 cm. Raceme 34- to
47-flowered, cylindrical. Pedicels 3.5–7 mm, erect
before flowering, then nodding or slightly deflexed.
Perianth 9–12 mm, tubular–campanulate, violet at
base and light violet above at anthesis, later brown-
ish, with greenish lobes with green stripes, ovate to
oblong. Anthers violet. Fruiting raceme cylindrical,
pedicels longer than capsule, horizontal or slightly
incurved. Valves of capsule globular–elliptic. Seeds
broadly ellipsoid to globular. – 2n=8 chromosomes.
Flowering: March to May.
Habitat: Damp meadows, open stony places, olive
groves and cultivated fields.
Distribution: Mediterranean element. South France,
Italy, Greece, Cyprus, Turkey, Egypt, western Syria,
Palestine. In Greece, it is distributed mainly in the
East Aegean Islands and also on the island of Kithira.
According to Feinbrun (1938–1940), this species is
considered as the most primitive among all extant
Bellevalia species and only presents slight morphologi-
cal variation throughout its geographical distribution.
3. Bellevalia romana (L.) Sweet, Hortus Brit.: 419
(1826) B. romana (L.) Reichenb., Fl. Germ.
Excurs.: 105 (1830) Hyacinthus romanus L.,
Mant. II.: 224 (1771). – Lectotype (Stearn, 1990:
216) (Italy) ‘Habitat Romae in cultis et hortis
sponte’, Herb. Linn. No. 438.10 (LINN, photo!)
(Fig. 3).
=Bellevalia operculata Lapeyr., Journ. Phys. 67:
426 (1808).
=Bellevalia appendiculata Lapeyr., Hist. Pl. Pyr.:
186 (1813).
=Scilla romana Ker Gawl., Bot. Mag. t. 939
=Bellevalia cyanoleuca St. Lag., Ann. Soc. Bot.
Lyon 7: 121 (1880).
Leaves 3–6, 25–50 cm ¥3–12 mm, longer than scape,
erect, linear, ascending, margins glabrous. Scapes
1–2, 24–36 cm. Raceme 15- to 30-flowered, oblong,
loose. Pedicels erecto-patent, (6.5–)7–12(–14) mm.
Perianth 7–9 mm, turbinate, whitish at anthesis,
sometimes light blue at the basis, later brownish,
with linear–oblong lobes equalling or longer than
tube. Anthers violet. Fruiting raceme cylindrical,
pedicels longer than capsule, erect–patulus. Valves of
capsule globular–elliptic. Seeds broadly ellipsoid to
globular. – 2n=8 chromosomes.
Flowering: March to April.
Habitat: Meadows, grassy places and cultivated
Distribution: Mediterranean element. In Greece it is
found in the Ionian Islands and the western and
southern continental parts (Ipiros, Sterea Hellas and
Feinbrun (1938–1940), in her monography of the
genus, considers previous reports from northern
Africa and Cyprus as erroneous. The species is
common and abundant in Italy, very rare in south-
western France and it extends eastwards to Greece
(Stearn, 1990), where it is also scarce and local.
4. Bellevalia ciliata (Cyr.) Nees., Gen. Pl. Fl. Germ.
4: t. 8 (1833–1837) Hyacinthus ciliatus Cyr., Pl.
Rar. Neap. 2: 23 (1788–92). – Type: Described from
Italy (Naples) (Fig. 4).
=Muscari ciliatum Ker Gawl., Bot. Reg. 5: t. 394
=Hyacinthus patulus Bertol., Misc. Bot. i: 21
Leaves 3–5, 32–37 cm ¥(9–)13–30 mm, shorter than
the scape, lanceolate, margins densely long–ciliate.
Scape 30–45 cm. Raceme 40- to 60(–70)-flowered,
conical. Pedicels patulous before anthesis, (15–)25–
64 mm, later recurved. Perianth 9–11 mm, campanu-
late, ovate, greenish or pale lilac before anthesis,
afterwards grey–brown, with greenish lobes 1/2 to 1/3
as long as tube, ovate. Anthers violet. Fruiting raceme
broadly conical, pedicels rigid, horizontal, 7–10 mm,
longer than capsule. Valves of capsule oblong–
triquetrate, emarginate, 14–15 mm. Seeds broadly
ellipsoid to globular. – 2n=8, 16 chromosomes.
Flowering: March to April.
Habitat: Open ground, fields and cultivated places.
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
Distribution: Widely distributed, mostly in the east
Mediterranean region. In Greece it is distributed
mostly in continental mainland (Macedonia, Thessa-
lia, Sterea Hellas and Peloponnisos).
The minor morphological differences between B.
ciliata and other related species, such as B. sarman-
tica (Pallas ex Georgi) Woronow and B. trojana
Feinbr., in combination with adjacent geographical
distribution, necessitate a more detailed study of the
species, based on living material, in order to resolve
the taxonomic status (Berg et al., 1989; Constantini-
dis, Kamari & Phitos, 1997).
5. Bellevalia edirnensis N. Özhatay & Mathew,
Botanical Journal of the Linnaean Society 107:
90 (1991). – Type: (European Turkey) ‘Edirne to
Uzunköprü, 1 km from Ibrik Tepe road junction,
250 m’, 30 April 1988, Güler Dalgiç 59752 (ISTE!)
(Fig. 8).
Leaves 3–4(–5), 30–42 cm ¥15–20 mm, longer than
the scape, very narrowly oblanceolate, undulate,
margins membranaceous, short ciliate. Scapes 1–2,
18–33 cm. Raceme 50- to 60-flowered, ellipsoid to
cylindrical. Pedicels erect at first, then arcuate,
8–15 mm. Perianth 8–11 mm, tubular–campanulate,
white with greenish base of tube, lobes slightly
shorter than the tube, elliptic–ovate. Anthers yellow.
Fruiting raceme cylindrical. Valves of capsule broadly
ovate, 9–10 mm. Seeds ovoid to ellipsoid. – 2n=24
Flowering: April to May.
Habitat: Edges of cultivated fields and meadows.
Distribution: European Turkey. NE Greece (Evros
province) [Fig. 9 (1)].
A new record for Greece of a species previously
considered to be a local of European Turkey. Among
the Bellevalia species occurring in Greece, B. edirn-
ensis is closely related to B. romana (Özhatay et al.,
1991b). The main morphological differences between
those two species are the short ciliate leaf margins,
the slightly shorter than the tube perianth lobes and
the yellow anthers of B. edirnensis, whereas in B.
romana the leaf margins are glabrous, the lobes are
as long as or longer than the tube and the anthers are
6. Bellevalia brevipedicellata Turrill, Kew Bull.
1940: 264 (1940). – Type: (Greece, SW Kriti) ‘above
Palaiochora at Kephala, stony limestone hills,
with Biarum davisii, 1 March 1940, Davis 1223K
(iso-E, photo!) (Fig. 5).
Leaves 2–3, 8–17 cm ¥10–20 mm, as long as the
scape, linear–lanceolate, undulate, margins glabrous.
Scapes 1–3, 9.5–16 cm. Raceme 15- to 26-flowered,
cylindrical, initially compact, later elongating.
Pedicels erecto-patent, 1–1.5 mm. Perianth 5–8 mm,
tubular to ± campanulate, white with lobes 1/2 to 1/3
as long as the tube, broadly ovate, flushed pink with
a green median line. Anthers violet. Fruiting raceme
cylindrical, pedicels shorter than capsule. Valves of
capsule obovoid to elliptic, 12–15 mm. Seeds broadly
ellipsoid to globular. – 2n=8 chromosomes.
Flowering: January to March.
Habitat: It grows in crevices and phrygana, on stony
limestone hills, from sea level up to 150 m.
Distribution: Endemic. Restricted to a limited
lowland area of southwestern Kriti [Fig. 10 (1)].
This taxon has been ranked as Vulnerable in the
Red Data Book of Rare and Threatened plants of
Greece (Turland, 1995). Its limited distribution area
is threatened by degradation or even destruction.
Furthermore, the potential value of the species in the
specialist horticultural market places it at the risk of
being overexploited through commercial bulb collec-
tion activities.
7. Bellevalia sitiaca Kypriotakis & Tzanoudakis,
Bot. Helv. 109: 85 (1999). – Type: (Greece, Kriti,
Prov. Lassithion) ‘prope vicum Sitia, in saxosis
calcareis promontorii Phaneromeni’, 2 April 1975,
Tzanoudakis 1814 (UPA!) (Fig. 6).
Leaves 3–5, 15–33 cm ¥5.5–15 mm, longer than the
scape, oblong, with visible parallel veins, margins
glabrous. Scapes 1–2, 12.5–27 cm. Raceme (13–)20- to
30-flowered, cylindrical. Pedicels 1 mm. Perianth 4.5–
7.5 mm, tubular cylindrical to tubular–campanulate,
white, with lobes 1/2 to 1/3 as long as the tube,
broadly ovate, flushed pink with a green median line.
Anthers violet. Fruiting raceme cylindrical, pedicels
shorter than capsule. Valves of capsule suborbicular,
10–15 mm. Seeds globular. – 2n=16 chromosomes.
Flowering: February to March.
Habitat: Rocky places and limestone cliff crevices at
low and moderate altitudes.
Distribution: Endemic. Restricted to eastern Kriti,
province of Sitia [Fig. 10 (2)].
Based on inflorescence and floral morphology, B.
sitiaca and B. brevipedicellata are obviously closely
related. However, their isolated geographical distri-
bution, as well as certain morphological differences
between the two species resulted in the description of
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
B. sitiaca as a new species (Kypriotakis & Tza-
noudakis, 1999). The leaves of B. sitiaca have visible
parallel veins, are thinner, longer and narrower than
those of B. brevipedicellata and the shape of the fruits
is suborbicular, not obovoid to elliptic as in B.
It is common in the Sitia province and prefers rocky
places. It grows in phrygana and limestone crevices,
where it occurs together with some characteristic taxa
of the Cretan flora, such as Viola scorpiuroides Cosson,
Scorzonera cretica Willd., Campanula pelviformis
Lam., Linum arboreum L., Ebenus cretica L., etc.
8. Bellevalia hyacinthoides (Bertol.) K. Persson &
Wendelbo, Bot. Notiser 132: 65 (1979) Strang-
weja hyacinthoides Bertol., Mem. Soc. Ital. Sci.
Modena 21,parte fisica: 3 (1837). – Type: ‘Floret in
horto Bot. Bononiensi Decembri decedente et toto
Januario, etiam sub dio. Patria hactenus ignota’
(BOLO? not seen) (Fig. 7).
=Hyacinthus spicatus Sm., Fl. Graec. Prodr. 1: 237
(1809) [non Moench, Meth.: 632 (1794)] Pusch-
kinia dubia Kunth, Enum. Pl. 4: 338 (1843) Bell-
evalia spicata Boiss., Diagn. Pl. Or. Nov., ser. 1, 7:
110 (1846) Foxia spicata (Boiss.) Parl., Nuov.
Figures 9–10. Distribution maps. Fig. 9, (1) B. edirnensis (the Greek population is indicated with blue colour, while the
Turkish populations with purple) and (2) B. hyacinthoides. Fig. 10, (1) B. brevipedicellata and (2) B. sitiaca.
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
Gen. Spec. Monocot.: 17 (1854) Strangweia
spicata (Boiss.) Boiss., Fl. Or. 5: 309 (1882). –
Syntypes: (Greece) ‘in insula Zacyntho’ Sibthorp et
‘in agro Argolico’ Sibthorp (OXF, not seen).
Leaves 3–6, (8.5–)13–30(–55) cm ¥(1–)1.5–7(–10) mm,
longer than the scape, basal, linear–lanceolate,
margins densely and shortly ciliate. Scapes 1–4,
(4.5–)5–18(–25) cm. Raceme 6–15-flowered, dense,
subspicate. Bracts lanceolate, deflexed, with a long
recurved basal lobe. Perianth 6–11 mm, campanulate,
pale blue, with recurved lobes 2 to 3 times as long as
tube. Filaments attached at base of lobes, with a long
tooth on each side. Anthers dark blue. Fruiting
raceme dense, subspicate. Valves of capsule broadly
elliptic to subreniform, 6–7 mm, emarginate-truncate
at apex. Seeds pyriform, testa smooth. – 2n=8+0–1B,
12 chromosomes.
Flowering: February to the end of March.
Habitat: It is usually found on dry stony slopes with
macchie and phrygana, edges of cultivated fields,
olive groves and abandoned fields, from sea level up
to c. 1000 m.
Distribution: Endemic. It is distributed in the
Kiklades Islands, the central and southern mainland
and the Ionian Islands [Fig. 9 (2)].
Persson & Wendelbo (1979), in their thorough study
of the species, comment extensively on its taxonomi-
cal status. Bellevalia hyacinthoides was previously
known as the Greek endemic monotypic genus
Strangweja, under the combination Strangweja
spicata (Sibth. & Sm.) Boiss., named in honour of
William Fox-Strangways, an English diplomat and
botanist. It differs from all other Bellevalia species in
having prominent teeth at the base of the filaments;
this was one of the main characters for its separation
at generic level by various authors (Mathew, 1994).
However, according to Persson & Wendelbo (1979),
the differences in filament morphology are not impor-
tant enough to support an independent genus. There-
fore, after re-examination of its morphology and
determining its basic karyotype formula, which was
found similar to that of most Bellevalia species, they
incorporated it in Bellevalia and described the section
Strangweja to accommodate its placement. Bellevalia
hyacintoides, besides its distinctly toothed filaments,
differs from other species of the genus by its dense
subspicate inflorescence, the comparatively large
bracts with a recurved basal lobe, the deeply lobed
perianth and the pyriform seeds.
Bellevalia hyacinthoides has an ornamental value,
equivalent to that of some Scilla and Muscari species
and it is certainly much more attractive than some
other Bellevalia species.
Although the genus Bellevalia is closely related to
Muscari,Hyacinthus and Hyacinthella, sharing with
them considerable morphological similarity, its karyo-
type is unique and clearly distinguishable. The ‘basic
karyotype’ of Bellevalia (von Bothmer & Wendelbo,
1981) consists of x=4, with long chromosomes (7.2–
15.8 mm). The karyotype formula can be described as
follows: one metacentric (m) chromosome, which is
the longest in the complement, one acrocentric (st)
chromosome being the second in size and two sub-
metacentric (sm) chromosomes. The similar size and
morphology of the two shorter chromosomes (both are
submetacentric) makes the distinction of the homo-
logues very difficult, despite the presence of satellites,
which may occur on both pairs. Small satellites,
sometimes conspicuous, appear among the species
often with variation in their number and position
among the species.
Polyploidy is also known to occur in Bellevalia and,
although diploid karyotypes are the most common
ones (Benzer, von Bothmer & Wendelbo, 1972; von
Bothmer & Wendelbo, 1981), tri-, tetra-, hexa- and
octoploids have been recorded up until now. The dif-
ferent ploidy levels found in Bellevalia, together
with the presence of B-chromosomes and other mor-
phological rearrangements suggest that the genus is
still under active chromosomal evolution (Johnson,
Most of the Greek Bellevalia populations karyologi-
cally examined revealed a diploid karyotype with
2n=2x=8. However, several polyploidy levels have
also been found in the Greek material, most of them
reported here for the first time.
1. Bellevalia dubia subsp. boissieri,2n=2x=8
(Fig. 11).
Karyotype formula: 2n=2x=2m-SAT +2st +4sm-
SAT =8.
All the populations of B. dubia subsp. boissieri
studied here, as well as those of B. romana and B.
trifoliata studied here, were found to be diploids.
Their karyotypes are mostly similar, differing prima-
rily in the position of the satellites and secondly in
their chromosome size and arm ratio.
In B. dubia subsp. Boissieri, the chromosome
number 2n=2x=8 has previously been reported in
material from Greece (von Bothmer & Benzer, 1973;
Bareka, Koutoula & Kamari, 2000; Kapasa et al.,
2001) and other countries (Garbari, 1968; Maggini,
1972; Maggini & de Dominicis, 1977; Özhatay,
Johnson & Mathew, 1991a; Johnson & Brandham,
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
1997). Additionally, a triploid chromosome number of
2n=3x=12 has been reported in material from Italy
(Musano & Maggini, 1976). In the chromosome
complement, six of the eight chromosomes bear small
spherical satellites on the shorter arms of the meta-
centric and on the long arms of the submetacentric
2. Bellevalia trifoliata,2n=2x=8 (Fig. 12).
Karyotype formula: 2n=2x=2m-SAT +2st +2sm
+2sm-SAT =8
The chromosome number of 2n=2x=8 and the
karyotype morphology of B. trifoliata has already
been given in material from Greece (von Bothmer &
Benzer, 1973) and from elsewhere (Feinbrun, 1938–
1940; Garbari & Tornadore, 1972; von Bothmer
& Wendelbo, 1981; Garbari, Giordani & Arnold,
1988; Özhatay et al., 1991b; Johnson & Brandham,
1997; Garbari, Giordani & Marchetti, 1999; Vogt
& Aparicio, 1999; Johnson, 2003). Therefore, B.
trifoliata appears to be ivariably diploid throughout
its geographical distribution. Its karyotype includes
two satellited chromosome pairs; the long metacentric
chromosomes bear small spherical satellites on their
shorter arm, while the short submetacentric chromo-
somes on their longer arm.
Figures 11–14. Microphotographs of mitotic metaphases and corresponding karyograms. Fig. 11, B. dubia subsp.
boissieri (2n=2x=8). Fig. 12, B. trifoliata (2n=2x=8). Fig. 13, B. romana (2n=2x=8). Fig. 14, B. edirnensis
(2n=6x=24). Scale bars, 10 mm.
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
3. Bellevalia romana (L.) Sweet, 2n=2x=8
(Fig. 13).
Karyotype formula: 2n=2x=2m-SAT +2st +2sm-
SAT +2sm =8.
The chromosome number of 2n=2x=8 has already
been given by several authors in material from various
countries (Delaunay, 1922; Darlington, 1926; De Mol,
1926; Feinbrun, 1938–1940; D’Amato, 1947; Jona,
1966; Garbari, 1968; Lovka et al., 1971; Maggini & de
Dominicis, 1977; Baldini, 1992, 1997). The chromo-
some number, as well as the karyotype morphology of
the Greek material, are given here for the first time
and are in accordance with previous reports. Bellevalia
romana, likewise B. trifoliata, has two satellited chro-
mosome pairs; the long metacentric chromosomes bear
small spherical satellites on their shorter arm. Addi-
tionally, satellites are found on the long arm of the
longest submetacentric chromosome pair.
4. Bellevalia ciliata,2n=2x=8 (Fig. 15) and
2n=4x=16 (Fig. 16).
Karyotype formulas: 2n=2x=2m-SAT +2st-SAT +
2sm +2sm-SAT =8.
2n=4x=4m-SAT +4st-SAT +4sm +4sm-SAT =16.
Most of the populations of B. ciliata studied here
are diploids, thus confirming previous references
(Feinbrun, 1938–1940; Constantinidis et al., 1997 and
references found therein) and possess six satellited
chromosomes. This is the only Greek taxon where
satellites are observed on the acrocentric chromo-
somes. Nevertheless, one population collected from
north-eastern Peloponnisos on serpentine substrate
was proved to be tetraploid with 2n=4x=16 chromo-
somes. This is the first report of polyploidy in this
5. Bellevalia edirnensis,2n=6x=24 (Fig. 14).
Karyotype formula: 2n=6x=6m +6st +12sm =
The newly discovered in Greece B. edirnensis has an
hexaploid karyotype, which is in accordance with the
chromosome number given by Özhatay et al. (1991b),
Özhatay et al. (1991a), Dane (1999) and Johnson
(2003) for the respective Turkish populations.
This is the first hexaploid Bellevalia recorded in
Greece. Several authors have indicated hexaploid
counts in other Bellevalia species; i.e. B. saviczii
Woronow in material from Afghanistan (Benzer et al.,
1972; Podlech & Bader, 1974; von Bothmer & Wen-
delbo, 1981) and Iran (von Bothmer & Wendelbo,
1981), B. glauca Kunth from Iran (von Bothmer &
Wendelbo, 1981), B. flexuosa Boiss. from Israel (von
Bothmer & Wendelbo, 1981) and B. alexandrina
Feinbr. from Egypt (Feinbrun, 1938–1940).
6. Bellevalia brevipedicellata,2n=2x=8
(Fig. 17).
Karyotype formula: 2n=2x=2m-SAT +2st +2sm-
SAT +2sm =8.
7. Bellevalia sitiaca,2n=4x=16 (Fig. 18).
Karyotype formula: 2n=4x=4m-SAT +4st +4sm-
SAT +4sm =16.
Besides the minor morphological differences found
in those Cretan endemics, B. brevipedicellata and B.
sitiaca also differ in their chromosome number. Bell-
evalia brevipedicellata is diploid with 2n=2x=8,
Figures 15–16. Microphotographs of mitotic metaphases and corresponding karyograms of B. ciliata. Fig. 15, 2n=2x=8.
Fig. 16, 2n=4x=16. Scale bars, 10 mm.
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
while B. sitiaca is tetraploid with 2n=4x=16 chro-
mosomes. The chromosome number of both taxa has
already been given by Tzanoudakis & Kypriotakis
(1987), Tzanoudakis et al. (1991) and Kypriotakis &
Tzanoudakis (1999).
The two species present more or less similar chro-
mosome morphology, even at the position of the sat-
ellites (Figs 17, 18). These are observed in both taxa
on the short arms of metacentric chromosomes, as
well as on the long arms of the longest in size sub-
metacentric chromosomes.
8. Bellevalia hyacinthoides,2n=2x=8+0–1B
(Fig. 19) and 2n=3x=12 (Fig. 20).
Karyotype formulas: 2n=2x=2m +2st +4sm =
2n=3x=3m +3st +6sm =12.
The 19 populations of the Greek endemic B. hya-
cinthoides, which have been cytologically studied by
us, cover all its geographical distribution and are
diploids, thus confirming previous counts from nine
populations given by Persson & Wendelbo (1979),
Speta (1979) and Bareka & Kamari (1999). Moreover,
in one population, located in Peloponnisos, close to
the city of Patras, one B-chromosome has been
observed (Phitos, 1988).
Figures 17–18. Microphotographs of mitotic metaphases and corresponding karyograms. Fig. 17, B. brevipedicellata
(2n=2x=8). Fig. 18, B. sitiaca (2n=4x=16). Scale bars, 10 mm.
Figures 19–20. Microphotographs of mitotic metaphases and corresponding karyograms of B. hyacinthoides. Fig. 19,
2n=2x=8. Fig. 20, 2n=3x=12. Scale bars, 10 mm.
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
The only exception to the diploid cytotype is one
population from Kefallinia (Ionian Islands), which we
found to be triploid, showing 2n=3x=12 chromo-
somes in all individuals examined. Triploidy is very
rare in the genus, as only one individual of B. dubia
from Italy (Musano & Maggini, 1976) and two indi-
viduals of B. sarmantica from the former Soviet
Union (Pogosyan, 1975; Pogosyan & Torosyan, 1983)
have been reported as such so far. No satellites have
been observed in the chromosome complement of B.
In this study, we have presented an extensive karyo-
systematic investigation of all Greek Bellevalia taxa
throughout their distribution range, based on numer-
ous populations. Following the discovery of the hexa-
ploid B. edirnensis in north-eastern Greece, the
number of species occurring in the country has now
increased from seven to eight.
Taking into account the infrageneric division of the
genus Bellevalia in sections by Feinbrun (1938–1940),
four of the Greek taxa, i.e. B. romana,B. edirnensis,B.
brevipedicellata and B. sitiaca, belong to section
Patens Feinbr., sharing significant morphological simi-
larity. Another two closely related taxa, which belong
to section Nutans Feinbr., are B. dubia and B. trifo-
liata. The most distant taxonomically taxa among the
Greek representatives of the genus are B. ciliata,
which belongs to section Conica Feinbr., and B. hya-
cinthoides of section Strangweja (Bertol.) K. Persson &
Wendelbo, previously considered as a separate genus.
The basic chromosome number of x=4, as well as
the karyotype morphology of all examined taxa, are in
accordance with the ‘basic’ Bellevalia karyotype. The
only difference detected among the species, in most
cases, is the position of satellites.
Bellevalia dubia subsp. boissieri,B. romana,B.
trifoliata and B. brevipedicellata are diploids, B.
sitiaca is always tetraploid and B. edirnensis always
hexaploid. New ploidy levels were found for two
diploid Bellevalia taxa in Greece: a tetraploid popu-
lation of B. ciliata growing on serpentine substrate
from Peloponnisos and a triploid population of B.
hyacinthoides from Kephallinia island.
The serpentine or ophiolithic rocks and soils have
long been known to support a specialized flora. The
tetraploid population on serpentine comes as no sur-
prise. Certain cases of differentiation and evolution
involving karyological data have also been investi-
gated. Recently, a karyological study of Greek serpen-
tine angiosperms made by Constantinidis, Bareka &
Kamari (2002) showed several changes in the karyo-
type morphology of plants growing on serpentine sub-
strate. The case of the tetraploid population of B.
ciliata is another example that provides evidence that
the ophiolithic substrate may be responsible for
karyological differentiation.
Although the genus Bellevalia is characterized by
particular karyotype morphology, it is noteworthy
that the position of the satellites may be variable but
stable intraspecifically in all populations examined.
B. dubia subsp. boissieri and B. ciliata have six
satellited chromosomes, with the later characterized
by the presence of satellites on the long arms of the
acrocentric chromosomes. B. romana and B. trifoliata
share considerable karyological similarity. The only
difference between these two species is observed at
the position of the satellites in the short submetacen-
tric chromosomes. B. romana bears satellites on the
longer submetacentric chromosome pair, while B. tri-
foliata on the shorter. B. edirnensis does not have
satellited chromosomes.
The two Cretan endemics, B. brevipedicellata and
B. sitiaca, despite having an isolated geographical
distribution and different ploidy levels (diploid and
tetraploid respectively), do have several morphologi-
cal similarities in combination, with a resemblance in
their chromosome complements concerning the posi-
tion of the satellites.
The species with the most evident morphological
differences and highest karyological variation, com-
pared with the other Bellevalia species, is B. hya-
cinthoides. Even at first glance, it does not appear to
belong to the genus and, although it has an almost
typical Bellevalia karyotype, the absence of satellites
in combination with the finding of one triploid popu-
lation and one with an accessory B-chromosome is
noteworthy indeed.
The presence of B-chromosomes is not very common
in the genus Bellevalia. However, the occurrence of
1–8 supernumerary chromosomes has been previously
reported by several authors (Zakhariyeva & Makush-
enko, 1969; Al-Mudarris, 1973; Johnson, 2003;
Gettner, 2005) in Asiatic species of the genus. The
only report of B-chromosome, originated from Euro-
pean material, has been reported by Phitos (1988) in
B. hyacinthoides.
To conclude, karyological characteristics as chromo-
some number, ploidy level, centromere position and
the number and location of satellites may be used in
elucidating taxonomical relationships of several plant
taxa (e.g. Sahin et al., 2006). However, because of
its relatively stable karyotype formula, the case of
Bellevalia is more complicated. Karyomorphometric
analysis does not provide clear results, as the only
difference among species is the number and morphol-
ogy of the satellited chromosomes. However, satellites
are sometimes difficult to locate because of their small
size and, therefore, a more detailed karyological work
on the genus appears necessary, which may include
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
fluorochrome banding, fluorescent in situ hybridiza-
tion (FISH), number of nucleoli and genome size.
Such a study is currently in progress by the authors.
The questions pertaining particularly to the taxo-
nomic classification of the Greek endemic Bellevalia
taxa require further study by also implementing dif-
ferent karyological and molecular techniques in order
to achieve a profound understanding of the taxonomy
and evolution of this attractive genus.
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The populations indicated by an asterisk have been
studied karyologically.
1. Bellevalia dubia subsp. boissieri
Kerkira Island: am Kastell, 19.III.1936, Ronniger
s.n. (5574 W); inter urbem Kerkira et vicum Kondokali,
in olivetis graminosis, 25.III.1972, Greuter 9773
(28970 ATH); Paleokastritsa und nähere Umgebung,
alt. 0–100 m, 2.IV.1985, Vitek CJ89 (2001-09974, 2001-
09976 W); Kerkira, Hlomos, olivenhain, 17.III.1990,
Borkowsky s.n. (UPA). Nomos Levkados: Levkas
Island: Kaligoni, 12.IV.1929, Just s.n. (12507 W); area
of the village Hortata, alt. 680–700 m, roadsides,
fallow fields and limestones, 16.IV.1970, Stamatiadou
8087 (6104 ATH); Mt Stavrota, summit Profitis Ilias,
alt. 1100–1150 m, stony, rocky calcareous ground,
22.IV.1977, Stamatiadou 19763 (38497 ATH); *from
the village of Nydri, on the way to the village of
Karyotes 1.5 km after the crossroads, 25.III.2005,
Trigou s.n. (UPA); *close to the village of Chortata,
25.III.2005, Trigou s.n. (UPA). Nomos Kefallinias:
Kefallinia Island: Ep. Samis: along the road from the
village Potamianata to Divarata, alt. 190–220 m, dry
stony places with olive trees, 15.IV.1967, Stamatiadou
70 (11216 ATH); Mons Kalon Oros, alt. 850 m, 38°19
N, 20°34E, 12.V.1973, Tzanoudakis 2414 (UPA); *ad
Ajia Varvara, 2 km ab urben Argostolion, ad viam,
38°09N, 20°30E, 21.II.1985, Phitos & Kamari 19409
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
(UPA); at the edges of the road toAgios Georgios castle,
alt. 180 m, 2.IV.2000, Phitos & Kamari 26477 (UPA,
MNHC-I); *Farsa, inside the district area, close to the
road, stony places, alt. 110 m, 19.II.2001, Katsouni 259
(UPA); *place named Avythos, Kalligata plain, olive
groves, alt. 90 m, 19.II.2001, Katsouni 256 (UPA);
Agios Konstantinos, alt. 7–8 m, uncultivated fields,
19.II.2001, Katsouni 258 (UPA); *Kardakata-Thinia,
above the village Kondogourata, alt. 230 m, unculti-
vated fields, 19.II.2001, Katsouni 257 (UPA); *Ep.
Kraneas, above the village of Davgata, cultivated
fields, alt. 220 m, 10.IV.2004, Phitos, Kamari & Kat-
souni 26949 (MNHC-I); *Ep. Palis: close to the village
Angona, place named Vrisi Petricaton, alt. 200 m, at
the edges of cultivated fields, 10.IV.2004, Phitos,
Kamari & Katsouni 26956 (UPA). Ithaki Island:
Place named Chani, 7.IV.2000, Katsouni s.n. (UPA);
Plateau called Anogi, 7.IV.2000, Katsouni s.n. (UPA).
Nomos Zakinthou: Zakinthos Island: Nordseite
des Vrachionas, 21.III.1936, Ronniger s.n. (5575 W);
obere Region des Skopos, 24.III.1936, Ronniger s.n.
(5606 W); summit area of Mt Skopos, between the
monastery of ‘Panajia Skopiotissa’ and Tourla, alt.
450–492 m, dry fallow fields and rocky shrubby slopes,
clayey-marly sandstones, gypsum and limestone,
13.IV.1971, Stamatiadou 11616 (14741 ATH); around
the monastery of Anafonitria, alt. 350 m, fallow or
cultivated fields, roadsides and open pinewood, lime-
stone, 14.IV.1971, Stamatiadou 11643 (13953 ATH);
village Orthonies, monastery of Spileotissis, alt.
400 m, fallow fields, S of monastery, stony ground with
phrygana and shrubs, 20.III.1976, Stamatiadou 18800
(26726 ATH). PELOPONNISOS: Nomos Achaias: *Ep.
Patron, above the village of Ano Kastritsi, edge of small
fields in clearings of macchie, alt. 600 m, 38°16N,
21°50E, 24.III.1990, Strid G901709 (ex cultis, UPA);
*Mt Erimanthos, on the way from Kalousi to Koumani
village, at the edges of the road, 1.II.2005, Bareka &
Lampropoulos Bel.71 (UPA); *from the village Stavro-
dromi to Roupakia, macchie, 22.IV.2005, Bareka &
Lampropoulos Bel.100 (UPA). Nomos Messinias: Ep.
Pilia, along earthen road between the villages Iamia
and Harokopio, near the settlement of Potamia, alt.
120–200 m, dry, stony roadbanks and fallow fields,
1.IV.1969, Stamatiadou 5113 (6107 ATH). Nomos
Lakonias: *Mt Parnon, summit Mavro Vouno, 4 km
from the village Metamorfosi to Richea, at the edges of
the road, limestone rocky places with Quercus coc-
cifera, Genista acanthoclada, Scabiosa hymettia etc.,
alt. 450 m, 36°49N, 22°55E, V.2003, Kalpoutzakis
s.n. (Bel.65 UPA). STEREA HELLAS: Nomos Attikis:
*Ep. Megaridos, Mt Kitheron, the chapel of Panagia
Gouras and the NW slopes of the summit Gennimata,
calcareous substrate sparcely covered with Phlomis
fruticosa and Quercus coccifera shrubs, alt. 600–
960 m, 38°11N, 23°22E, 23.IV.1995, Constantinidis
5364 (UPA); Egina Island: Ajios Nektarios monastery,
18.II.1968, Post 909 (5396 ATH). Nomos Pireos:
Kithira Island: Cerigo, 1894, Makowsky (W). Nomos
Fokidos: Ep. Parnassidos, 3–4 km WNW from the
village Antikira, along the communal road to the
village Dhesfina, alt. 400 m, dry–rocky hillsides with
phrygana and Euphorbia dendroides shrubs above the
road, limestone, among rocks by a sheepfold,
30.III.1968, Stamatiadou 1852 (3865 ATH); W of
Dhelfi, by the road to Itea, alt. 500–550 m, stony fallow
fields with scattered shrubs of Quercus coccifera, lime-
stone, 26.III.1972, Stamatiadou 14376 (20470 ATH).
Nomos Viotias: Ep. Levadhias, 1 km before the
village Aliartos, by the road from Thiva to Levadhia,
alt. 100–130 m, dry–rocky hillsides with phrygana,
limestone, 30.III.1968, Stamatiadou 1798 (6761 ATH);
Ep. Levadhias, 4 km outside of Levadhia, along the
road to Arahova, shrubby hillsides and streams above
the road, damp places, schistose ground, 30.III.1968,
Stamatiadou 1814 (6105 ATH); SW of the town of
Levadhia, Ajios Ierousalim, alt. 600–700 m, wood of
Pinus,Cupressus and Laurus, scarce, on rocks,
13.III.1972, Petamidis 851 (20588 ATH); coast of Dhis-
tomon, NW of the hamlet ‘Aspra Spitia’, alt. 50–150 m,
stony ground with olive trees in lower zone and rocky
limestone slopes with evergreen shrubs (Pistactia
lentiscus,Quercus coccifera,Phillyrea,Euphorbia
dendroides,Phlomis fruticosa etc.), 25.III.1972,
Stamatiadou 14272 (20366 ATH). Nomos Etolias-
Akarnanias: Ep. Mesolongiou, c. 3kmSEofAno
Mousoura, somewhat damp, shady places in Quercus
coccifera scrub by a small stream, alt. 400–500 m,
38°28N, 21°26E, 3.V.1997, Strid S975702 (ex cultis
UPA); mons Koutsilaris, alt. 0–200 m, 2.IV.1981,
Christodoulakis et al. 619 (UPA); NW of Aetos village,
garigue, dry place, alt. c. 550 m, 38°43N, 21°03E,
5.III.1997, Vlachos 2/31 (UPA); *2 km after the village
Aggelokastro, at the edges of the road, wet meadows,
25.III.2004, Bareka & Lampropoulos Bel.85 (UPA);
*5 km after Lessini village outside the monastery, wet
meadows, 25.III.2004, Bareka & Lampropoulos Bel.87
(UPA); *1 km after Astakos, by the road, olive groves,
25.III.2004, Bareka & Lampropoulos Bel.89 (UPA).
IPIROS: Nomos Prevezis: Ep. Nikopoleos-Pargas,
Mt Zalogon, between the village of Arhea Kassopi
and Zalogon monument, alt. 600–750 m, stony rocky
calcareous ground, 30.IV.1978, Stamatiadou 20862
(40643 ATH). MACEDONIA: Nomos Pierias: Ep.
Pierias: gravelly flats between Litochoron and the sea,
alt. 50–100 m, 40°08N, 22°33E, 23.III.1987, Chris-
tensen, Møkker & Strid sub Strid no 25403 (UPA).
ITALY:Sicily, Palermo, April, Gussone s.n. (NAP,
photo!); in pascuis montosis supra Bocca di Falco,
20.III.1855, E. & A. Huet du Pavillon s.n. (W).
CROATIA:Dalmatia, Lesina, 1989, Botteri s.n. (W).
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
2. Bellevalia trifoliata
GREECE:STEREA HELLAS: Nomos Attikis: In insula
Aegina, 18-26.IV.1881, Heldreich s.n. (UPA). Nomos
Pireos: Kithira Island: *1 km SE of Kalamos,
recently burnt area in guarigue and small olive grove,
alt. 100 m, 36°09N, 23°02E, 22.III.1998, Strid &
Tan S982401 (ex cultis UPA). EAST AEGEAN
ISLANDS: Nomos Lesvou: Lesvos Island: close to
Skala Pamfilon, locality named Nisselia, 8.IV.1993,
Bazos IB801 (ATHU); ibid., 1.IV.1998, Bazos IB3047
(ATHU). Nomos Chiou: Chios Island: village Neo-
horion, grassy places and cultivated fields around the
cemetery ‘Ajios Prokopios’, alt. 80 m, sandy–clayey
ground, 7.III.1969, Stamatiadou 4441 (6109 ATH).
Nomos Dodekanisou: Island of Ro: bay of
Frangolimnionas, alt. 30 m, stony to rocky calcareous
ground with phrygana and shrubs, 26.III.1974,
Stamatiadou 17652 (34348 ATH). Kastellorizo
(Megisti) Island: the plain (ipsipedho) of Vikla, alt.
230 m, compact terra rossa soil, 5.IV.1973, Stamatia-
dou 16566 (29740 ATH); the plain of Dhexamenes
(water reservoirs), alt. 40–60 m, shady places in olive
grove, 8.IV.1973, Stamatiadou 16647bis (29882 ATH);
the cape E of the Harbour, between Conaki and the
Lycian tomb, alt. 5–30 m, shady, rocky calcareous
slopes above the sea, 22.III.1974, Stamatiadou 17473
(34163 ATH); the plain (ipsipedho) of Ajios Jeorgios
tou Vounou, in fallow fields SW of monastery, alt.
150 m, compact terra rossa soil with phrygana,
23.III.1974, Stamatiadou 17528 (34221 ATH). Rodos
Island: in campis insulae Rhodi, III.1845, Heldreich
462 (W); valley of Petaloudhes, alt. 130–150 m, shady
areas by the river, 3.IV.1973, Stamatiadou 16466
(29640 ATH).
CYPRUS:District Cyrenia, Pelea Vrysi – Nordhang,
alt. 900 m, northern range, 25.III.1973, Bauer &
Spitzenberger 23A (23178 W); *Akamas, area called
Rigenas castle, Cistus-Juniperus growth, 24.XI.2001,
Ch. Georgiadis s.n. (UPA); *Chrysochou, phrygana,
25.XI.2001, Ch. Georgiadis s.n. (UPA).
TURKEY:A2(E), Belgrad ormani (Istanbul) (çayir),
29.IV.1956, A. & E. Baytop 4472 (ISTE); A2(E),
Hadimköy (Istanbul), 23.IV.1961, A. & E. Baytop 6337
(ISTE); A2(E). Terkos Köyü (Istanbul) – nemli çayir –,
1.V.1967, A. & T. Baytop 10930 (ISTE); A2(E), Yildiz
Parki (Istanbul), 4.IV.1972, E. Tuzlaci 21380 (ISTE);
A2(E) Halkali (Istanbul) tren istasyonu, çayirlar,
23.IV.1971, N. & E. Özhatay & G. Ertem 19360
(ISTE); A2(E), Uskumruköy (Istanbul) – kumlu
tepenin kars¸isindaki sirtlar, 29.IV.1971, G. Ertem, F.
Öktem, N. & E. Özhatay 19424 (ISTE); A2(E), Terkos
köyü çayiri (Istanbul), 3.V.1971, A. Baytop, G. Ertem,
N. Özhatay 19453 (ISTE); A2(E), Çatalca’ya 4 km
(Istanbul), 18.V.1975, N. & E Özhatay 31637 (ISTE);
A2(E), Çatalca, Hadimköy, Tugay, Servi korusu
(Istanbul), alt. 60 m, 28.IV.2002, Ï. Genç 1228, 82102
(ISTE); A2(E), Çatalca, Izzettin, Çadirtepe mevkii
(Istanbul), alt. 25 m, 17.IV.2003, I. Genç 1584, 82365
(ISTE); A2(E), Istanbul, Yildiz parki (Istanbul), A.
Baytop 55030 (ISTE); A2(A), Yalova üstü, 3.V.1950, T.
Baytop & A. Berk 2470 (ISTE); B1, Bornova, Manisa
yolu (Izmir), 6.V.1961, T. Baytop 6462 (ISTE); B1,
Ayvalik, Hakkibey Yarimadasi güneydog˘usu, Timar-
hane Burnu civari (Balikesir), alt. 10 m, 15.IV.1998,
K. Alpinar 74959 (ISTE); B1, Ayvalik, Küçük Maden
Adasi (Balikesir), alt. 20 m, 16.IV.1998, K. Alpinar
75042 (ISTE); C3, Kemer, Olimpos Adrasan arasi,
Çayalani köyün –den 1 km (Antalya) terkedilmis
tarla–, 23.III.1973, H. Pes¸men & K. Güner (HP 4235)
52621 (ISTE); C4, Antalya, Nerencine bahçelerinde,
25.II.1966, A. & T. Baytop, N. Tanker & E. Sezik 8554
(ISTE); C6 Hatay, 8 km SE Samandag, Flussufer,
2.IV.1985, Sorger 85-11-14 (1991-03155 W);
LEBANON/SYRIE:Beryti, III.1894, Renzevalle s.n.
(1996 04353 W);
FRANCE:Toulon, IV.1857, Chambeiron s.n. (8277 W);
ibid., 20.III.1869, Chambeiron s.n. (W); Var: La
Garde, près Toulon, dans les champs de la plaine,
12.IV.1888, Albert s.n. (8257 W).
ITALY:Bordighera, Liguria, 2.IV.1890, Bicknell s.n.
(10994, 10751 W); ibid., 14.IV.1891, Bicknell s.n.
(11181 W); ibid., IV.1892, Bicknell s.n. (1283 W); ibid.,
24.IX.1896, Bicknell s.n. (10751 W).
3. Bellevalia romana
Kerkira Island: Corfu, 1894, Makowsky s.n. (8226
W); inter urbem Kerkira et vicum Kondokali, in olive-
tis graminosis, 25.III.1972, W. Greuter 9774 (28938
ATH); near Kerkira town, place named Kanoni, lower
Kanoni road, wet fields, 29.II.1972, Baxter 65 (23368
ATH); Insel Korfu, Paleokastritsa und nähere
Umgebung, alt. 0–100 m, 2.IV.1985, Vitek CJ89
(2001-09976 W); *Ep. Kerkiras: near the village of
Kanakades, cultivated fields and meadows on deep
clay, seasonally wet, alt. 80 m. 39°39N, 19°46E,
8.IV.1997, Strid S970201 (ex cultis UPA). Nomos
Levkados: Levkas Island: plain of Vasiliki, road-
sides, meadows and wet marshy brooksides near the
sea, sandy-clayey soil, 15.IV.1970, Stamatiadou 8029
(6108 ATH); Agios Nikitas, alt. 0–10 m, 26.III.2005,
Trigou s.n. (UPA). Nomos Kefallinias: Kefallinia
Island: c. 1–2 km S Agrapidies, gegen den Giupari, c.
800–900 m, Felsentriften, Schutt, Kalk, 12.V.1980, E.
& F. Krendl s.n. (2001-13468 W). PELOPONNISOS:
Nomos Lakonias: *Monemvasia, outside the castle
at the edges of the road, IV.2003, Sagona s.n. (UPA).
STEREA HELLAS: Nomos Etolias-Akarnanias:
*2 km S before Astakos, in petrosis et saxosis calc.,
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
25.III.1995, Phitos, Kamari, Greuter & Zimmer Bel.21
(UPA); Mt Boumistos, 1 km west of Komboti village,
along the road leading to deserted Komboti village,
streams and pastures, alt. c. 500 m, 38°45N, 21°04
E, 12.IV.1998, Vlachos 40/610 (UPA). IPIROS: Nomos
Thesprotias: Ep. Thiamidos, 1 km SE of the village
of Plataria, seasonally wet meadows, alt. 5–10 m.
39°27N, 20°16E, 16.IV.1989, Strid & al. G890201
(ex cultis, UPA); Nomos Prevezis: Ep. Nikopoleos-
Pargas, 1 km SW of Nikopolis, alt. 0–5 m, clayey
cultivated field, 39°01N, 20°44E, 29.III.1987, Chris-
tensen, Møller & Strid Strid25473 (UPA).
LIBYA:(B. romana var. mauritanica) Cirenaica, Tra
el-Gubba e u el-Atrun Argub Dasc, 14.III.1934, Pam-
panini & Richi-Sermolli 1514 (19402 W);
CROATIA:Dalmatia: Litoral Plflanze, in kleinen wäs-
serigen Gräben zwischen Aeckern in Castel-Abadessa
bei Spalato, Petter 204 (4123, 17315, 21940 W).
ITALY:Habitat Romae in cultis et hortis sponte, Herb.
Linn. No. 438.10 (LINN, photo!); in cultis agrillosis
planitiei Catania, 25.III.1856, E. & A. Huet du Pavil-
lon 201 (W); Rome, villa Medici, IV.1865, Parseval-
Grandmaison s.n. (W); in pratis agri florentini,
28.III.1872, Levier s.n. (14931 W); copiosa in agris et
incultis circa Florentiam, III.1873, Levier s.n. (12279
& 14930 W); in herbosis incultis agri Florentini,
28.IV.1874, Levier s.n. (99481, 10979 W); Boboli
garten in Florenz, 1.IV.1875, Riechter s.n. (6632,
15871 W); prati humidi nei dintorni di Modena,
IV-V.1881, Mori s.n. (14003, 14004 W); in pratis
humidis Florentiae, IV.1883, Bergeesti s.n. (41584 W);
in agri Florentino, IV.1883 Martelli s.n. (6631 W);
agro Florentino, IV.1887, Martelli s.n. (5349 W); prati
dintorni di Modena, IV.1888, Mori s.n. (14932 W); nei
prati d’ intorno a Modena, IV.1890, Mori s.n. (11744
W); dintorni di Modena, IV.1891, Mori s.n. (15868 W);
Modena, 27.IV.1936, Cortesi III854 (6977 W); Roma,
Sibthorp (W); Calabria, in ducatu parmensi (624,
276712 W); Ravena, am Friendhof von Comacchio,
8.IV.1950, Höpflinger (888 W).
FRANCE:Corsica: prairies de l’étang de Biguglia,
2-16.IV, Mabille 1867 (155031 W); Var: La Crau
d’Hyéres, prés et champs humides, 28.III.1889, Albert
2367 (538 W); Hérault: prairies de Coursand,
21.V.1892, Mouret 3126 (4828 W).
4. Bellevalia ciliata
Ermionidos: Didima, just S of the village, cultivated
fields and olive groves with many perennial weeds, alt.
120 m, 37°27N, 23°10E, 1.IV.1998, Strid & Tan
S989402 (ex cultis UPA); Between Palea and Nea
Epidavros, at the edges of cultivated fields, 12.IV.2000,
Phitos, Kamari & Constnantinidis K407 (UPA).
Nomos Arkadias: *Ep. Kinourias, Mt Parnon, from
Agios Andreas village to Prastos village, 3 km after
Dipotamo, plateau Chourio, cultivated fields, alt. c.
600 m, 7.IV.2002, Kalpoutzakis s.n. (UPA). STEREA
HELLAS: Nomos Attikis: in agris inter Oropo et
Chalcidem, 4.1884, Heldreich s.n. (W); in arvis rarior,
legimus pr. Liosia ad radices m. Parthenis, Fl.
29.III.1889, fruit 26.IV.1889 Heldreich 1082 (14933,
3120 W; UPA); pr. Liosia ad radices m. Parthenis,
Heldreich 1082 (3119 W). Nomos Viotias: *Mt Kith-
eron, c. 1.5 km south-west of the village of Kaparelli,
edges of cultivated fields, c. 450 m, 38°13N, 23°12E,
17.IV.1993, Constantinidis 22984 (UPA); *c. 5km
north of Ritsona area, cultivated land, c. 240–270 m,
38°25N, 23°31E, 20.III.1994, Constantinidis 4322
(UPA); *Ep. Thivon, c. 2.5 km ENE from the village of
Dafni, on the way to the village of Dafnoula, cultivated
and abandoned fields, c. 400 m, 38°15N, 23°26E,
16.IV.1994, Constantinidis 4436 (UPA). THESSALIA:
Nomos Larissis: Larissa, in agris incultis versus
Nehali, 14.V.1927, Rechinger fil. 1132 (17652 W).
MACEDONIA: Nomos Thessalonikis: in collinis ad
Thessalonicam, V.1903, Adamovic´ s.n. (W).
5. Bellevalia edirnensis
GREECE:THRAKI: Nomos Evrou: *Ep. Didimoti-
chou: c. 3 km NW of the village Koufovouno, at the
edges of cultivated fields, 3.V.2004, Bareka & Lam-
propoulos Bel.76 (UPA).
TURKEY:A1 (E), Edirne: Edirne to Uzunköprü, 1 km
from Ibrik Tepe road junction, alt. 250 m, 30.IV.1988,
G. Dalgiç 59752 (ISTE, Typus!); A1 (E), Merkez
mezarlig˘i altlari, Fen Fakültesi bahçesi (Edirne),
25.IV.1989, G. & A. Dalgiç 60218 (ISTE).
6. Bellevalia brevipedicellata
GREECE: Kriti Island: Nomos Chanion: above
Palaiochora at Kephala, stony limestone hills, with
Biarum davisii, 1.III.1940, P. Davis 1223 (E, Isotypus,
photo!); Chrysoskalitisa, 28.I.1991, Kypriotakis s.n.
(Isotypus, UPA); prope Chrysoskalitisa, 8.II.1991,
Kypriotakis s.n. (Isotypus, UPA); *by the monastery of
Chrysoskalitisa, alt. 0–25 m, limestone, 5.IV.2002,
Bareka & Tzanoudakis Bel.41 (UPA).
7. Bellevalia sitiaca
GREECE: Kriti Island: Nomos Lasithiou: Eparhia
Sitias: prope vicum Sitia, in saxosis calcareis prom-
ontorii Phaneromeni, 2.IV.1975, Tzanoudakis 1814
(UPA, Typus); near the village Palaekastro, 1.II.1991,
Kypriotakis 17/45 (UPA); ibid., 8.II.1991, Kypriotakis
13/4 (UPA); the gorge near the village Adravastos,
19.II.1991, Kypriotakis 15/55 (UPA); ibid.,
12.III.1992, Tzanoudakis s.n. (UPA); *road to Mari-
dati beach, E of road from Vai to Palekastro, alt.
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
30 m, 35°13N, 26°15E, 2.IV.2003, Bareka &
Turland 0958a (UPA); *3 km after the village of Agia
Fotia on the way to the village of Palaekastro, alt.
8–9 m, rocky cliffs, phrygana, 2.IV.2003, Bareka &
Turland Bel.61 (UPA); *2.5 km from Vai to Palaekas-
tro, after the crossroads to Maridati beach, alt. 30 m,
garigue, 2.IV.2003, Bareka & Turland Bel.60 (UPA);
*on the way to Maridati beach, alt. 3–5 m, garigue,
2.IV.2003, Bareka & Turland Bel.59 (UPA); *on the
way from Zakros to Xerokambos village, alt. 80 m,
limestone crevises, 2.IV.2003, Bareka & Turland
Bel.62 (UPA); *from Xerokambos to Chametoulo
village, Chametoulo gorge, alt. 40–50 m, rock cre-
vises, 2.IV.2003, Bareka & Turland Bel.58 (UPA).
8. Bellevalia hyacinthoides
GREECE:THESSALIA: Nomos Pierias: Prope Lito-
choro Thessaliae in regione inferiori Olympi,
3-4.1870, Krüper s.n. (10241 ATH). Nomos Larissis:
*Along the road, 1.5 km before the village Omolio, alt.
50–80 m, 39°53N, 22°42E, 27.IV.1999, Bareka &
Barekas Bel.37 (UPA). STEREA HELLAS: Nomos
Etolias-Akarnanias: NW of Aetos village, garigue,
dry place, alt. c. 550 m, 38°43N, 21°03E, 5.III.1997,
Vlachos 2/47 (UPA); E of Panagoula village, stream
Tourkos, phrygana, limestone cliffs, alt. c. 700 m,
38°43N, 21°00E, 12.IV.1997, Vlachos 6/63 (UPA);
NE of Panagoula village around Panagia chapel,
garigue, dry place, alt. c. 550 m, 38°43N, 21°00E,
18.V.1997, Vlachos 6/64 (UPA); *1.5 km before the
village Aggelokastro, outside the castle of Kira Rini,
rocky hillsides, 25.III.2004, Bareka & Lampropoulos
Bel.86 (UPA); *1 km after Astakos, by the road, stony
places, 25.III.2004, Bareka & Lampropoulos Bel.88
(UPA). Nomos Fokidos: Ep. Parnassidos: 6 km from
Itea along main coastal road to Navpaktos, secondary
shrubland with Phlomis fruticosa, alt. 20 m, 38°26N,
22°24E, 17.II.1989, Strid 28005 (UPA). Nomos
Viotias: Oberhalb Arachowa am Parnass, alt.
c. 1050 m, 20.IV.1933, Ronninger s.n. (5573 W); *Ep.
Thivon, the bay of Domvrena, E parts of Paralia Agios
Nikolaos, c. 3.9 km NW Aliki, slopes facing the seam
phrygana and Juniperus phoenicea shrubs, eroded
ophiolitic substrate, alt. c. 3–20 m, 38°12N, 23°01E,
8.III.1999, Constantinidis 8324 (UPA). Nomos
Attikis: Pr. Clucine collib. marit., 24.II.1844, Hel-
dreich s.n. (10236 ATHU); ad radices m. Hymmeti
pr. Bari, 29.I.1874, Heldreich s.n. (10242 ATHU); in
collibus aridis regionis littoralis prope Trachones,
25.I.1875, Heldreich s.n. (10237 ATHU, 12283 W);
Habit prope Eleusinam (rara), fl. Jan.-Febr.,
10-22.II.1850, Orphanides 141 (10238 ATHU);
Eleusis, IV.1844, Heldreich s.n. (W). PELOPONNISOS:
Nomos Achaias: *Prov. Achaia, infra pagum Kastrit-
sion, ad Panepistimioupolis, 38°17N, 21°47E,
I.1970, Phitos 2823 (UPA); infra pagum Kastritsion,
ad Panepistimioupolis, 38°17N, 21°47E, 13.II.1975,
Artelari 11 (UPA); prope pagum Kataraktis, in petro-
sis, alt. c. 500 m, 38°05N, 21°50E, 18.II.1989,
Phitos & Kamari 20308 (UPA); *3 km before the
village Ellinikon, at the edges of cultivated fields,
1.II.2005, Bareka & Lampropoulos Bel.72 (UPA);.
Nomos Arkadias: *Ep. Kinourias, Zavitsi Mt, 2.8 km
from Xiropigado to Kato Vervena village, at the edges
of a stream, alt. 50–60 m, 37°27N, 22°44E,
9.III.2003, Kalpoutzakis 1245 (UPA). Nomos Argoli-
dos: prope Naupliam Peloponnesi, Heldr. 1845,
Orphanides s.n. (10240 ATHU); in collibus agri
Argolici, primo vere, Sartori 6 (W); in collibus agri
Argolici, Sartori 70 (276730 W). Nomos Lakonias:
*Mt Parnon, summit Koulochera, 800 m from the
crossroads to the church of Profitis Ilias, limestone
rocky places with Quercus coccifera, Genista acantho-
clada, Euphorbia rigida etc., alt. 450 m, 36°49N,
22°55E, V.2003, Kalpountzakis s.n. (Bel.64 UPA).
IONIAN ISLANDS: Nomos Levkados: Levkas
Island: Megas Oros, alt. 1016 m, 16.IV.1929, Just s.n.
(12394 W); *on the way to the village Syvota, olive
groves, alt. 30–70 m, 9.X.1999, Bareka & Lampropou-
los Bel.20 (UPA); *from the village of Englouvi, on the
way to the village of Chortata, rocky places with
macchie, 25.III.2005, Trigou s.n. (UPA); in ins.
Leucade, Mazziarii s.n. (10235 ATH). Nomos Kefall-
inias: Kefallinia Island: *mons Aenos in silva
Abietis cephalonicae, alt. 1100 m, 38°02N, 20°37E,
31.III.1971, Phitos & Kamari 11111 (UPA); ad
Castrum pagi Assos, 38°22N, 20°32E, 30.I.1970,
Phitos & Kamari 10789 (UPA); close to the village of
Agios Athanasios, 1993, Katsouni s.n. (MNHC-I);
*place named Trapezaki at the edges of the road,
27.I.2001, Katsouni s.n. (UPA); *Ajia Varvara, place
named Diatsentia, 22.I.2001, Katsouni s.n. (UPA); *at
the entrance of Cyclopean walls, 26.I.2001, Katsouni
s.n. (UPA); *Davgata, regional road to Langada, close
to the district area, fields, alt. 280 m, 26.I.2001,
Katsouni s.n. (UPA, MNHC-I); *at the bridge of
Ajia Varvara, close to Argostoli, calcareous rocks,
alt. 80 m, 27.I.2001, Katsouni s.n.( MNHC-I); *Ep.
Pallis, Patrikata Erissou, inside the build-up area,
alt. 250 m., 9.III.2006, Katsouni s.n. (UPA); *Vary
Erissou, at the edges of the road, phrygana,
9.III.2006, Katsouni s.n. (UPA); area named Zamoli,
regional road to Ichthiotrofia, olive groves,
10.III.2006, Katsouni s.n. (UPA); *above the village of
Chavriata, olive groves, alt. 200 m, 10.III.2006, Kat-
souni s.n. (UPA); *Ep. Karaneas, Kardakata Thinias,
at the edges of olive groves, alt. 190 m, 9.III.2006,
Katsouni s.n. (UPA). Nomos Zakinthou: Zakinthos
Island: nordseite des Vrachionas, 21.III.1936, Ronni-
ger s.n. (5607 W). KYKLADES ISLANDS: Nomos
Kikladon: Folegandros Island: In insula Pholegan-
dro, II.1863, Veneris s.n. (10239 ATH).
© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 723–739
... In Greece, eight taxa of the genus have been recorded to date (Bareka 2008; Bareka et al. 2008Bareka et al. , 2012 (2n ¼ 4x ¼ 16), both belonging to B. sect. Bellevalia and restricted to the island of Kriti (Crete). ...
... Karyology: The karyological study of Bellevalia juliana revealed a hexaploid karyotype with 2n ¼ 6x ¼ 24 chromosomes. The morphology of the chromosome complement follows the basic Bellevalia karyotype (Bareka 2008;Bareka et al. 2012;, while at least two satellite submetacentric chromosomes are observed. The karyotype formula consists of 2n ¼ 6x ¼ 6m þ 6st þ 10sm þ 2sm-SAT ¼ 24 chromosomes, varying in size from 16.5 to 8.26 mm (Figure 2). ...
... Kunth (Zakhariyeva & Makushenko 1969;) and B. sarmatica (Johnson & Brandham 1997). So far, hexaploid populations have been evidenced in several species (Table II), only one species of which, Bellevalia edirnensis, from European Turkey and NE Greece, seems to be exclusively hexaploid (Ö zhatay et al. 1991a, 1991bBareka 2008;Bareka et al. 2008Bareka et al. , 2012. ...
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A new species, Bellevalia juliana Bareka, Turland & Kamari (Asparagaceae) is described from E Kriti (Greece) and compared with other related taxa from Greece and the Mediterranean area. It is a member of B. sect. Bellevalia and a karyological study revealed a hexaploid karyotype with 2n = 6x = 24 chromosomes, illustrated here. The conservation status of the new species is assessed as Vulnerable (VU), according to IUCN criteria.
... On the basis of morphological criteria, they have been traditionally linked together (Garbari and Greuter 1970) and their close relationship was supported by molecular phylogeny, placing them in the same clade (Pfosser and Speta 1999). Moreover, the geographical range of both genera covers the same areas from the western Mediterranean region (Morocco, Algeria) eastwards throughout Europe and southwestern Asia (Johnson 2003, Nersesian 2001, Bareka et al. 2008, Jafari et al. 2008, Jafari 2012a, 2012b, Borzatti Von Loewenstern et al. 2013, Demirci et al. 2013). However, from the karyological point of view, Bellevalia and Muscari, differ significantly from each other. ...
... According to Brullo et al. (2009), the Tunisian species show a close relationship with B. pelagica C. Brullo, Brullo & Pasta, 2009 also tetraploid, and endemic to Lampione islet (Sicily). Cytogenetic studies (Bareka et al. 2008(Bareka et al. , 2012 and phylogenetic analysis (Borzatti Von Loewenstern et al. 2013), performed on populations occurring in Greece and Italy respectively, highlighted the diversity in Bellevalia and raised questions about the taxonomic relationships and the origin of polyploids. ...
... Usually, in the genus Bellevalia, the karyotypes show satellites on either the first, the second or the third pair of chromosomes (Bothmer and Wendelbo 1981, Bareka et al. 2008Bareka et al. , 2012. Our tetraploid B. mauritanica from Stidia shows a similar chromosome arrangement and bears one pair of satellites on the first metacentric pair. ...
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Karyological investigations were carried out on four species of Bellevalia Lapeyrouse, 1808 and Muscari Miller, 1758 (Asparagaceae) sampled in contrasting bioclimatic conditions of Algeria. The endemic Bellevalia mauritanica Pomel, 1874 was found to have a tetraploid cytotype 2n = 4x = 16 and an octoploid 2n = 8x = 32 which is a new report. The chromosome number 2n = 2x = 18 in Muscari comosum (Linnaeus, 1753) Miller, 1768 and Muscari maritimum Desfontaines, 1798 was in conformity with earlier reports. The latter species reveals a lesser bimodality of the karyotype. Within Muscari neglectum Gussone ex Tenore, 1842 pentaploid (2n = 5x = 45), hexaploid (2n = 6x = 54) and very rare octoploid cytotype (2n = 8x = 72) have been reported in Algeria. Principal component analysis performed on basis of karyotype parameters, showed a segregation of the different cytotypes. This study provides new karyological information, which is discussed in a taxonomic context.
... The genus Bellevalia Lapeyrouse (1808: 425) belongs to the former Hyacinthaceae, particularly to the tribe Hyacintheae, and is represented by about 65 taxa globally. It is distributed over the Mediterranean region from Morocco and Algeria eastwards to the Caucasus and Iran (Bareka et al. 2008, Borzatti von Loewenstern et al. 2013. The last revision of the genus dates back to Feinbrun's monograph (1940), which included 44 species, eight of which distributed along the Turkish border. ...
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Bellevalia bayburtensis (Asparagaceae) is described and illustrated as a new species from Bayburt Province in northeastern Anatolia, Turkey. Diagnostic morphological characters, a full description, detailed illustrations, and a distribution map are provided. It is morphologically similar to B. rixii and B. paradoxa, but clearly differs from these species by the leaf shape, the bud and flower colour.
... Bellevalia has been deeply studied from the karyological point of view (i.e. Feinbrun 1938;Chiarugi 1949;Bentzer et al. 1972;Pogosyan 1975;Bothmer & Wendelbo 1981;Garbari 1981;Johnson 2003; Bareka et al. 2008Bareka et al. , 2012Azizi et al. 2016), due to its peculiar karyotype and to the role of polyploidy driving speciation in some taxa. Currently, within the genus, polyploidy has been documented in ca. ...
In the south-central Mediterranean four tetraploid species of Bellevalia occur: B. dolichophylla, B. galitensis, B. mauritanica, and B. pelagica. Another group of plants, morphologically similar to B. dolichophylla, has been recently recovered in Zembra Island (Tunisia). A phylogenetic reconstruction involving all these tetraploid taxa was performed using both plastidial and nuclear markers (trnL-trnF and ITS, respectively). For all these taxa, an allopolyploid origin involving B. romana and B. dubia is supported. Regarding plants from Zembra, they may fall within the variability of B. dolichophylla.
... Generally, karyological characteristics, as chromosome number, ploidy level, centromere position, and the number and location of satellites and secondary constrictions, can be used in elucidating taxonomical relationships of several plant taxa (Bareka et al. 2008, 2012 see for references). Although, karyomorphometrics is able to provide more information about the studied taxa, the conclusions can be used only as additional evidences to the primary hypothesis. ...
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Fritillaria Linnaeus, 1753 (Liliaceae) is a genus of geophytes, represented in Greece by 29 taxa. Most of the Greek species are endemic to the country and/or threatened. Although their classical cytotaxonomic studies have already been presented, no karyomorphometric analysis has ever been given. In the present study, the cytological results of Fritillariamontana Hoppe ex W.D.J. Koch, 1832 group, which includes Fritillariaepirotica Turrill ex Rix, 1975 and Fritillariamontana are statistically evaluated for the first time. Further indices about interchromosomal and intrachromosomal asymmetry are given. A new population of Fritillariaepirotica is also investigated, while for Fritillariamontana, a diploid individual was found in a known as triploid population. Paired t-tests and PCoA analysis have been applied to compare the two species.
... Scilloideae, tribe Hyacintheae (APGIII 2009;Chase et al. 2009). This genus is distributed over the Mediterranean region from Morocco and Algeria eastwards to the Caucasus and Iran (Özhatay et al. 1991, Bareka et al. 2008, Jafari & Maassoumi 2008, Borzatti von Loewenstern et al. 2013, Karabacak et al. 2014 and 74 species of this genus are listed in The World Checklist of Seed Plants (Govaerts 1996). ...
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Bellevalia behcetii (Asparagaceae) is described and illustrated as a new species from Şırnak province in South Eastern Anatolia (Turkey). It is morphologically similar to B. paradoxa but easily differs in several morphological characters. Diagnostic morphological characters, a full description and detailed illustrations of the new species are provided. A quantitative morphometric analysis was carried out to check the diagnostic characters of Bellevalia behcetii with respect to morphologically related species. The somatic chromosome number was determined as 2n = 2x = 8 in B. behcetii. In addition, the ecology and phenology of the new species, as well as its etymology and the pollen characteristics, are presented and discussed. The conservation status of B. behcetii was assessed according to IUCN protocol. A distribution map of B. behcetii and related species is also presented.
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This is the twelfth of a series of miscellaneous contributions, by various authors, where hitherto unpublished data relevant to both the Med-Checklist and the Euro+Med (or Sisyphus) projects are presented. This instalment deals with the families Asparagaceae (incl. Hyacinthaceae), Boraginaceae, Cactaceae, Caryophyllaceae, Chenopodiaceae, Compositae, Crassulaceae, Euphorbiaceae, Gramineae, Haloragaceae, Iridaceae, Labiatae, Leguminosae, Malvaceae, Orchidaceae, Orobanchaceae, Plumbaginaceae, Polygonaceae, Rosaceae, Scrophulariaceae (incl. Buddlejaceae), Solanaceae and Umbelliferae. It includes new country and area records and taxonomic and distributional considerations for taxa in Abutilon, Aegilops, Amelanchier, Andryala, Aruncus, Asparagus, Bellevalia, Brugmansia, Buglossoides, Bupleurum, Cortaderia, Crassula, Datura, Dysphania, Euphorbia, Fallopia, Iris, Lycianthes, Myriophyllum, Nicodemia, Onobrychis, Ophrys, Opuntia, Orobanche, Phelipanche, Plumbago, Salvia, Silene, Stellaria and Wisteria, and new combinations in Amelanchier and Phelipanche.
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Özet: Bu çalışmada, endemik ve tehlike altında olan Bellevalia edirnensis'in mevcut yayılışı, popülasyonlarının son durumu ve habitatını tehdit eden unsurlar araştırılmıştır. Elde edilen bulgular doğrultusunda CR kategorisindeki türün habitatında bü-yük problemler (aşırı otlatma, tarımsal faaliyetler, yapılaşma, su ve toprak kirliliği) olduğu gözlenmiştir. Bu problemlere in situ ve ex situ çözüm önerileri sunulmuş ve tartışılmıştır. En kısa sürede etkili bir koruma stratejisi oluşturulmadığı takdirde bu türün yakın zamanda doğada yok olacağı gerçeği öne sürülmüştür. Abstract: In this study, current distribution, population status and the threats on the habitat of the endemic and endangered species Bellevalia edirnensis were investigated. The findings indicated that overgrazing, agricultural activities, construction and water and soil pollution were the major threatened factors on the habitat of this critically endangered (CR) species. In situ and ex situ conservation strategies were presented and discussed. It is suggested that B. edirnensis is prone to extinction in nature in near future unless an effectiveconservation strategy is established as soon as possible.
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Bellevalia Lapeyrouse (1808: 425) is a genus comprising about 65 species and subspecies distributed over the Mediterranean region (Bareka et al. 2008, Jafari & Maassoumi 2008, Borzatti von Loewenstern et al. 2013). Wendelbo (1984) reported 18 taxa, seven of which endemic to the country. In further studies, Wendelbo (1985) reduced Bellevalia latifolia Feinbrun (1940: 369) to a synonym of B. olivieri (Baker 1874: 8) Wendelbo (1985: 120). Since then, three species were published by Ozhatay (2000), while Johnson (2003) reduced B. pycnantha (Koch 1849: 255) Losinskaja (1935: 310) to a synonym of B. paradoxa (Fischer & Meyer 1835: 30) Boissier (1882: 308). In recent years, four more species, B. leucantha Persson (2006: 253), B. malatyaensis Uzunh. & H.Duman in Uzunhisarcikli et al. (2013: 652), B. chrisii Yildirim & B.Şahin in Yildirim et al. (2015: in press) and B. pseudolongipes Karabacak & Yildirim in Karabacak et al. (2014: 210) were described. In this study, we present a further new species, Bellevalia koyuncui , endemic to Turkey.
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This is the twenty-three of a series of reports of chromosomes numbers from Mediterranean area, peri-Alpine communities and the Atlantic Islands, in English or French language. It comprises contributions on 56 taxa: Anthriscus, Bupleurum, Dichoropetalum, Eryngium, Ferula, Ferulago, Lagoecia, Oenanthe, Prangos, Scaligeria, Seseli and Torilis from Turkey by Ju. V. Shner, T. V. Alexeeva, M. G. Pimenov & E. V. Kljuykov (Nos 1768-1783); Astrantia, Bupleurum, Daucus, Dichoropetalum, Eryngium, Heracleum, Laserpitium, Melanoselinum, Oreoselinum, Pimpinella, Pteroselinum and Ridolfia from Former Jugoslavia (Slovenia), Morocco and Portugal by J. Shner & M. Pimenov (1784-1798); Arum, Biarum and Eminium from Turkey by E. Akalin, S. Demirci & E. Kaya (1799-1804); Colchicum from Turkey by G. E. Genç, N. Özhatay & E. Kaya (1805-1808); Crocus and Galanthus from Turkey by S. Yüzbaşioǧlu, S. Demirci & E. Kaya (1809-1812); Pilosella from Italy by E. Di Gristina, G. Domina & A. Geraci (1813-1814); Narcissus from Sicily by A. Troia, A. M. Orlando & R. M. Baldini (1815-1816); Allium, Cerastium, Cochicum, Fritillaria, Narcissus and Thymus from Greece, Kepfallinia by S. Samaropoulou, P. Bareka & G. Kamari (1817-1823).
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Constantinidis, Th., Kamari, G. & Phitos, D.: A cytological study of 28 phanerogams from the mountains of SE Sterea Ellas, Greece. — Willdenowia 27: 121–142. 1997. — ISSN 0511-9618. 28 phanerogams of various families, from the mountains of Gerania, Pateras, Kitheron, Pastra and Elikon (SE Sterea Ellas, Greece) are cytologically studied. The chromosome numbers of 13 taxa (Aristolochia microstoma, Asperula baenitzii, A. pulvinaris, A. rigidula, Centaurea subsericans, Conium divaricatum, Johrenia distans, Peucedanum vittijugum subsp. vittijugum, Scorzonera serpentinica, Thlaspi pindicum, Thymus parnassicus, Th. teucrioides subsp. candilicus, and Verbascum boissieri) are presented for the first time. In addition, Greek populations of 11 taxa are cytologically examined for the first time. Mitotic metaphase photomicrographs and/or karyograms are presented for the majority of taxa studied. Brief comments are given on the karyotype morphology, cytogeography and relationships of selected taxa.
Bellevalia sitiaca from Sitia, the easternmost district of Crete, is described as a new species to science. It is closely related to B. brevipedicellata Turrill, endemic to western Crete, but it differs in both morphology and karyology. Specimens of the new taxon in fruiting stage have been collected several years ago but were misidentified either as B. brevipedicellata or as Muscari macrocarpum Sweet. It would seem that all previous records of the latter from the Cretan area are based on specimens belonging to the new species now described; thus the occurrence of M. macrocarpum on the island of Crete remains questionable.
An interesting new finding of Bellevalia trifoliata in Tuscany is recorded. This diploid (2n=8) rare Mediterranean geophyte grows in a sunny slope near Ripa (Seravezza commune), Versilia area (Province of Lucca) at about 100 m a.s.l. on acid soil ("verrucano"). Its presence may be related to the migration of the termophilous floristic elements on the Tyrrhenian coasts after the last Würmian glaciation phases.
[French] Après avoir passé en revue et illustré par des exemples les principales causes d'erreur dans les comptages de chromosomes, l'auteur donne quelques recommandations pratiques pour les éviter le plus possible. Il montre que chez des polyploïdes élevés, il est parfois extrêmement difficile (sinon impossible) de savoir si le nombre zygotique est constant ou non dans une population. Il pense que la variation intraspécifique par polyploïdie ou dysploïdie précède assez souvent la variation interspécifique et il montre combien il faut être prudent dans l'établissement du nombre de base x d'un groupe donné. /// After having reviewed and illustrated with examples the chief reasons of mistakes in chromosome counting, the author gives practical recommandations to shun them as often as possible. He shows that with high polyploids, it is sometimes very difficult to know if the zygotic number is or is not constant in a population. He thinks that the intraspecific variation by polyploidy or dysploidy often precedes the interspecific variation and he shows how careful one must be in the establishment of the basic number of a studied group.