53 PHYTOLOGIA BALCANICA 19 (1): 53 66, Sofia, 2013

The genus Crocus (Iridaceae) in Greece: some noteworthy

floristic records and karyotypes
Theophanis Karamplianis
1
, Spyros Tsiftsis
2
& Theophanis Constantinidis
1
1
Department of Ecology & Systematics, Faculty of Biology, National &
Kapodistrian University of Athens, Panepistimiopolis, GR-157 03 Athens, Greece,
e-mail: [email protected], e-mail: [email protected] (corresponding author)
2
Department of Botany, School of Biology, Aristotle University of Thessaloniki,
GR-541 24 Thessaloniki, Greece, e-mail: [email protected]
Received: January 17, 2013 Accepted: March 25, 2013
Abstract. Since its first record in 1846, Crocus reticulatus is rediscovered in northern Greece. C. biflorus subsp.
alexandri, a rare species in Greece, is confirmed from east Macedonia. The populations of C. pallasii subsp.
pallasii are not confined to the East Aegean Islands only but extend to Macedonia, Thessaly and Sterea Ellas
of the Greek mainland as well, southwards to Mt. Imittos. Descriptions of these three taxa are provided
and their apparent relationships discussed. Furthermore, photographs of metaphase plates, karyotypes and
idiograms of all three taxa are reported: Crocus reticulatus has 2n = 12, C. biflorus subsp. alexandri 2n = 8 and
C. pallasii subsp. pallasii either 2n = 14 or 2n = 16.
Key words: chorology, Crocus, chromosome numbers, floristics, Greece, idiograms, new records.
Introduction
The genus Crocus L. (Iridaceae) comprises c. 140 taxa
(species and subspecies) in its whole distribution area,
including the cultivated species C. sativus L. (Mathew
1982, 1983, 1988, 2000a, 2000b; Rukns 2010). The
genus is confined to the Old World and extends lon-
gitudinally from the Iberian Peninsula to west China
and latitudinally from Poland and the Caucasus re-
gion to the northern parts of the Arabian Peninsula
(Mathew 1982; Rukns 2010).
Crocus exhibits a pronounced infraspecific vari-
ation with respect to morphology and chromosome
numbers, particularly in the eastern parts of its distri-
bution (Balkan Peninsula and Anatolia). According to
Maw (1886) and Feinbrun (1958) the genus appears
to have its origin in this latter area. Opposite theories
argue that the origin of Crocus should be placed in
northern Africa, particularly Cyrenaica where C. bou-
losii Greuter is found (Greuter 1968), and the Iberian
Peninsula where some primitive species (C. carpeta-
nus Boiss. & Reuter, C. nevadensis Amo) occur. These
species exhibit a semi-terete cross section of leaf and
share habitat similarities with the related genus Romu-
lea (Goldblatt 1971; Mathew & Brighton 1975). Ac-
cording to recent phylogenetic studies (Petersen &
al. 2008; Seberg & Petersen 2009), the most probable
place of origin for Crocus is North Africa and the Ibe-
rian Peninsula, although only a small number of taxa
are indigenous in these areas.
On Greek territory, 34 Crocus taxa are found
(Mathew 1982, 1983, 1988, 2000a, 2000b; Papanico-
laou & Zacharof 1980; Phitos & Kamari 1983; Rukns
2010, Karamplianis & al. 2011), including C. sativus.
This number accounts for about 25 % of all Crocus
taxa known so far. An important proportion, 14 spe-
cies or subspecies, should be considered as Greek en-
demics. Crocus boryi J. Gay and C. hadriaticus Herbert
54 Karamplianis, T. & al. Floristic records and karyotypes of Greek Crocus
subsp. hadriaticus, often considered as Greek endem-
ics, are shared also with Albania (e.g. Shuka 2008) and
should be considered regional or Balkan endemics.
Both have been collected by us a several scores or sev-
eral hundred of meters away of the Greek-Albanian
border, on Greek territory, and their population obvi-
ously continues further north within Albania.
This report deals with 3 Crocus taxa inadequately
known in Greece, i.e. Crocus reticulatus Steven ex Ad-
am, C. biflorus Mill subsp. alexandri (Nici ex Velen.)
B. Mathew and C. pallasii Goldb. subsp. pallasii.
Material and methods
All investigations and descriptions are based on Greek
plant material, either dry (see Appendix), deposited at
the Herbarium of National and Kapodistrian Univer-
sity of Athens (ATHU; Thiers, continuously updated),
or alive. The terms Nomos and Eparchia used in the
Appendix refer to administrative divisions of Greece.
Living material is being kept in cultivation at a
small experimental garden in the Faculty of Biology,
Athens University. Although cultivation is generally
successful, taxa of high mountains do not flower reg-
ularly at the low altitude where the collection is kept,
and slowly deteriorate. The classification and nomen-
clature mostly follows Mathew (1982). Description of
capsule and seed is based on collected samples with
additional information by Skourtis & Thanos (2012).
The karyological study was carried out in the De-
partment of Ecology & Systematics, Athens Univer-
sity, and is based on material collected from natural
populations (see Table 1 and Appendix). Root tips
were pretreated in an aqueous solution of 8-hydroxy-
quinoline (0.003 % w/v) for 44 h at room tempera-
ture, or less often in a saturated aqueous solution of
-bromonaphthalene for 24 h at 4 C. The materi-
al was then fixed in Carnoy solution (3:1 v/v absolute
ethanol: glacial acetic acid) for at least 24 hrs at 4 C
and stored in a solution of 70 % ethanol at -20 C. For
chromosome preparations the root tips were hydro-
lyzed in 1N HCl for 11 min at 60 C, stained with Feul-
gens stain for 34 hours and macerated in 45 % (v/v)
glacial acetic acid. Metaphases were observed using a
Zeiss Axio Imager A1 microscope and captured with
an AxioCam MR3. Selected photos were measured
using Image J v. 1.44p (Rasband, continuously updat-
ed). Chromosome terminology mostly follows Levan
& al. (1964). Karyotype asymmetry is calculated using
Stebbinss asymmetry index (SA; Stebbins 1971), total
form percentage index (TF %; Huziwara 1962), ratio of
long arm per total length of chromosome set (Ask %;
Arano 1963), and mean length of short arms per mean
length of long arms 100 index (Syi %; Greilhuber &
Speta 1976). These indexes are among the most accu-
rate in describing karyotype asymmetry, according to
Zuo & Yuan (2011).
Provenance of material, karyotype formulas and
asymmetry indexes are summarized in Table 1.
Results and discussion
An investigation of Falakron Mountain (east Mace-
donia, north-eastern Greece) in March 2009 by Spy-
ros Tsiftsis resulted in the discovery of two interest-
ing Crocus taxa. The same mountain was visited again
by S. Tsiftsis and Theophanis Karamplianis on March
30, 2010 and both alive and pressed plant specimens
were collected. The taxa were determined as Crocus
reticulatus and C. biflorus subsp. alexandri and were
seen growing in separate populations or occasionaly
mixed together in the same locality. Crocus reticula-
tus was rediscovered in Greece, after its first record by
Grisebach (1846) on Mt. Chortiatis (in pratis m. Kor-
thiat). Its occurrence on this latter mountain, howev-
er, was not reconfirmed in a thorough investigation by
Karagiannakidou & Raus (1996). The subpopulations
on Mt. Falakron consist of numerous individuals in-
habiting at least three different localities.
The second taxon, Crocus biflorus subsp. alexan-
dri, is rare and scattered in Greece. It has earlier been
reported from Mt Falakron by Petersen & al. (2008),
while Rukns (2010) mentions it from central Thra-
ce without providing any further details. Its possible
occurrence on Mt Athos (Agion Oros) is further dis-
cussed below.
Crocus pallasii subsp. pallasii was so far known in
Greece from the phytogeographical region of the East
Aegean Islands (EAe, Mathew 1982; Christodoula-
kis 1986; Panitsa 1997; Bazos 2005). Mathew (1999)
also gave North Greece as a distribution area of the
subspecies, but without referring to any precise lo-
calities. This subspecies has now been found in sev-
eral places of the Greek mainland during field work
conducted between 2008 and 2012. Its southernmost
known locality is on Mt Imittos, close to Athens. Fur-
55 Phytol. Balcan. 19(1) Sofia 2013
ther collections were made close to the city of Elasso-
na in Thessaly (Central Greece, autumn 2008) and on
Mt Vertiskos (central Macedonia, see Appendix). Two
more collections come from Mt Menikion (autumn of
2009 and 2010) and a hill east of Drama (east Macedo-
nia, November 2012).
Crocus reticulatus Steven ex Adam, Beitr. Naturk.
(Weber & Mohr) I: 45. (Fig. 1)
Description: Corm subglobose to ovoid, flattened at
the base, 1520 mm in diameter, covered by coarse-
ly reticulate-fibrous tunics with fibres up to 0.5 mm
wide, splitting into a coriaceous disc with radial fibres
at base and forming a neck with flaccid and strong fi-
bres upwards. Cataphylls 34, papery, white, occa-
sionally green at the apex. Leaves 45, synanthous,
reaching perianth throat at flowering time, 5 cm
(0.5)1(1.6) mm and up to 20 cm long at maturity,
glabrous, sparsely papillose at the margins and along
the convex keel beneath, with a white stripe above
up to half the lamina width, and two channels along-
side keel below, each with two prominent ridges. Pro-
phyll absent. Bract and bracteole membranous, white,
equal to subequal in length, the bracteole narrow-
er than bract. Flowers 12; tepals unequal, 1738
413 mm, acute to obtuse, inner ones broader than
outer, white or pale-lilac, outer ones white, lilac, sil-
very or buff, with 3 wide stripes and 23 secondary
longitudinal violet veins at outer surface, throat white
or often pale-yellow, glabrous to slightly papillose.
Filaments 57 mm, white, glabrous; anthers yellow,
1012(15) mm long, about twice the length of fila-
ments. Style somewhat exceeding stamens, branched
at around the middle of anthers into 3 yellow to deep
orange branches, expanded and papillose at apex, 14
20(-25) mm long. Capsule 1518 710(14) mm, el-
lipsoid, with three chambers each 78 mm wide. Seeds
brownish, occasionally pale-yellow (immature?) 3.0
3.5 1.52.5 mm with a prominent raphe beginning
about halfway the underside and ending at the apex of
the caruncle. Caruncle not so distinct and flattened,
reaching 0.5 1 mm, paler than the rest of seed, cha-
laza yellowish, rough, with ridges lacking any pattern,
testa brownish, somewhat smooth and papillose.
Distribution and ecology: Crocus reticulatus is dis-
tributed from Northeast Italy, Croatia and Slovenia
Fig. 1. Flowering plants
of Crocus reticulatus (a);
a group of plants in their
habitat (b); details of a fow-
ering individual (c) and the
structure of the outer corm
tunics (d). (Photographs by
S. Tsifsis).
56 Karamplianis, T. & al. Floristic records and karyotypes of Greek Crocus
(Ranelovi & al. 1990; Rpert 2000 onwards), Serbia
and Bosnia & Herzegovina (Ranelovi & al. 1990) east-
wards to Hungary (Ranelovi & al. 2007), Crimea, the
Caucasus area (Mathew 1982; Ranelovi & al. 1990;
Rpert 2000 onwards) and Turkey (Mathew 1984). In
the neighbourhood of Greece, it has been found in F. Y.
R. Macedonija (Ranelovi & al. 1990; Ranelovi & al.
2007), Bulgaria (Ranelovi & al. 1990; Ranelovi &
al. 2007; Vladimirov 2007; Fgras & al. 2010; Assyov
& Petrova 2012) and Romania (Ranelovi & al. 1990;
Fgras & al. 2010).
Three subpopulations of Crocus reticulatus have
been seen on Mt Falakron (Fig. 4), between 600 m and
1470 m a.s.l. (see Appendix). Their preferred habitat is
the clearings in Quercus, Fagus and Pinus forest, with
Juniperus communis subsp. nana, J. oxycedrus, Pulsa-
tilla halleri subsp. rhodopaea, Galanthus elwesii subsp.
elwesii, Crocus biflorus subsp. alexandri, and Primula
vulgaris. Plants were also found growing at the mar-
gins of melting snow patches in subalpine grasslands,
together with Crocus flavus, C. olivieri subsp. olivieri,
Sesleria sp., Rosa sp., and Scilla bifolia s.l. The geologi-
cal substrate of these localities consists mostly of mar-
bles and the soil is fertile due to the high content of
rhizomull humus. The plants flower from mid-Febru-
ary to mid-May, depending on how mild or harsh the
weather is.
The population on Mt Falakron consists of at least
3000 individuals and is not under any immediate
threat at present. It should be stressed however, that
massive collection, particularly for horticultural use,
should be strongly discouraged.
Taxonomic relationships: Crocus hittiticus Baytop &
B. Mathew, with a very narrow distribution in the Ci-
lician Taurus of South Turkey, is the closest relative of
C. reticulatus. The former species is often considered
a subspecies of C. reticulatus under combination C.
reticulatus subsp. hittiticus (Baytop & B. Mathew) B.
Mathew, according to Mathew (1982), but the differ-
ences between the two taxa are significant. C. reticu-
latus has very coarsely-reticulate corm tunics, with fi-
bres reaching a width of c. 0.5 mm, the leaves have two
channels alongside a keel underneath, each with two
prominent ridges, the anthers are yellow before dehis-
cence and the outer tepals are narrower than the inner
ones, a rare character for Crocus. In contrast, the corm
tunics of C. hittiticus have finer fibres of interwoven
texture, the underside of leaves has a keel but no ridg-
es along lateral channels (Kandemir 2010), the anthers
are maroon to blackish, with yellow pollen, and the
outer tepals vary from narrower to broader, as com-
pared to the inner ones. Furthermore, the two taxa do
not appear in close phylogenetic positions (Petersen
et al. 2008; Seberg & Petersen 2009). Nevertheless, the
members of the Reticulati series (where C. reticulatus
and C. hittiticus belong), Speciosi and Biflori, are in-
termixed in the known phylogenetic trees, forming a
moderately supported clade that indicates close and
complex relationships. On the basis of the above data,
we prefer to regard the two taxa as distinct at species
level until their affinities are fully elucidated.
On grounds of tunic structure, Crocus reticulatus
shares the characteristic wide fibres with C. cancella-
tus Herbert subsp. mazziaricus (Herbert) B. Mathew
of the same series.
Crocus biflorus Miller subsp. alexandri (Nii ex
Velen.) B. Mathew, Crocus: 85 (1982) (Fig. 2)
Description: Corm flattened-globose, 715 mm in di-
ameter, tunics membranous or coriaceous, forming
entire or toothed rings at base, with an indistinct corm
neck of triangular teeth mostly not exceeding 5 mm
in length. Cataphylls 35, papery, usually yellowish
or brownish, often distinctly speckled brownish-red.
Leaves 36, synanthous, up to 1.5 mm wide, with a
white stripe less than a quarter of laminas width and
without ribs on the underside. Prophyll absent. Bract
and bracteole present, subequal to unequal. Perianth
segments 2035 717 mm, obtuse or rounded but
often subacute, white inside, the exterior surface of
inner segments stained violet-blue at base, the exte-
rior of outer segments entirely stained violet-blue, oc-
casionally densely feathered, except for the narrow
white margin. Throat of perianth lacking yellow col-
ouration, glabrous. Filaments 37 mm long, white,
glabrous or seldom finely papillose; anthers 814 mm
long, yellow. Style divided into three yellow to red-
dish-orange slender branches, in some plants some-
what expanded at apex. Capsule ellipsoid, 1015 mm
long, carried on a short pedicel just above ground level
at maturity. Seeds subglobose and flattened, 2.53.0
1.52.0 mm, reddish-brown with an indistinct raphe.
Raphe somehow brighter than seed colour, beginning
at the middle of the underside and running through
seed surface to the apex of the caruncle, where strong-
ly attached. Caruncle prominent, 0.51.0 mm long, up
57 Phytol. Balcan. 19(1) Sofia 2013
to 1.0 mm wide. Texture with smooth testa and sparse
papillae on the raphe.
Distribution and ecology. Crocus biflorus subsp. al-
exandri is distributed in Bulgaria (Assyov & Petrova
2012), Serbia and F. Y. R. Macedonija (Ranelovi &
al. 1990). A specimen from Mt Athos (Agion Oros)
collected on February 6
th
1914 by Hartmann (B, pho-
to!) and seen by Bornmller in 1914 and B. Mathew
in 1973 may be the first record of this subspecies in
Greece. However, any recent confirmation from the
same area would be much welcome, since identifica-
tion of old dry specimens belonging to the critical C.
biflorus group is not always a straightforward task.
Recent Greek records of the same subspecies are
provided by Petersen & al. (2008) and Rukns (2010)
from Mt Falakron and Thrace, respectively, without
further details. All records appear on the map of Fig. 4.
On Mt Falakron, a large and continuous population
of Crocus biflorus subsp. alexandri has been observed,
distributed along the southern slopes of the moun-
tain and particularly above the village of Pyrgi, at c.
7001500 m. In some localities it grows sympatrically
with other Crocus species. The plants are found either
in openings amongst scattered Fagus and Pinus trees
where the soil is particularly rich in organic matter,
or in overgrazed grasslands and pseudomaquis forma-
tions consisting mainly of scattered Quercus coccifera
individuals. However, the most common habitat for
C. biflorus subsp. alexandri is the clearings in Fagus
and Pinus forests and the subalpine grasslands, where
it flowers near melting snow patches. This last hab-
itat is often shared with C. reticulatus, C. flavus and
C. oli vieri subsp. olivieri. The geological substrate con-
sists of marble or limestone. Habitat details are much
like those of C. reticulatus reported before. The sub-
species flowers from the end of February to the begin-
ning of April or possibly later, depending on the melt-
ing of snow cover.
Crocus biflorus subsp. alexandri is not particularly
rare on Mt Falakron. According to a preliminary esti-
mation, more than 5000 mature individuals are grow-
ing in situ. The population is distant from any settle-
ment or anthropogenic activities and does not seem
to be under any direct threat at present. It is not nega-
tively affected by overgrazing either. However, its nar-
row distribution in Greece, compared to other species
of the genus, may call for regular monitoring in the
future.
Taxonomic comments. Crocus biflorus subsp. alexan-
dri is easily distinguished from its Greek allies. It differs
from the typical subspecies, C. biflorus subsp. biflorus,
in the characteristic flower coloration of outer tepals:
they are stained with violet in a feathery way, leaving
only a small white margin untouched, a unique feature
among the Greek Crocus species. In subsp. biflorus, the
tepals are more or less similar to the other three Greek
subspecies, i.e. subsp. melantherus (Boiss.& Orph.) B.
Mathew, subsp. stridii (Pap. & Zach.) B. Mathew and
subsp. nubigena (Herbert) B. Mathew, with yellow
throat and three broad longitudinal stripes on the exte-
Fi g. 2. Fl ower i ng
plants of Crocus bi-
florus subsp. alexan-
dri (a); lateral view of
fower (b) and detail of
corm (c). (Photographs
by S. Tsifsis).
58 Karamplianis, T. & al. Floristic records and karyotypes of Greek Crocus
rior of the outer tepals that branch into secondary stri-
ations towards the tepal margin. The last three subspe-
cies also have in common the blackish-maroon anthers
(occasionally yellow in subsp. stridii), best observed
before dehiscence. In addition to the above, C. biflo-
rus subsp. melantherus is an autumn-flowered plant
distributed in South Greece. C. biflorus subsp. nubige-
na has two pairs of ribs on the underside of leaf (Kan-
demir 2011) and is found in the East Aegean Islands
(Raus 1983; Christodoulakis 1986, 1996; Bazos 2005).
C. biflorus subsp. stridii has papillose to sparsely pubes-
cent throat and filaments, and ciliate (occasionally on-
ly sparse) leaves with a distinctly broader white stripe
that reaches 2/3 of leaf width (Papanicolaou & Zacha-
rof 1980). It occurs on Mt Chortiatis, east of Thessalon-
iki and the area around Xanthi (Mathew 1995).
Perhaps the closest relative of Crocus biflorus sub-
sp. alexandri is C. biflorus subsp. weldenii (Hoppe &
Furnr.) B. Mathew, which differs in having ribs or
ridges on the underside of leaf. This latter subspecies
is known from the Balkans, i.e. north Albania (Rakaj
2009), Croatia (evrnja & Vladovi 2005), Montene-
gro, F. Y. R. Macedonija, Serbia, Bosnia & Herzegovi-
na, Slovenia, and also Northeastern Italy (Ranelovi
& al. 1990). It has not been recorded in Greece so far.
The group of Crocus biflorus is by far the most di-
verse and complex in the genus and still needs care-
ful investigations based on field work, dry specimens
and cultivated material coupled with experimental
approaches. Its members have been treated in vari-
ous ways by relevant botanists, particularly in region-
al Floras. Their patterns of variation and distribution
need to be further studied.
Crocus pallasii Goldb. subsp. pallasii, Mem. Soc.
Nat. Moscou 5: 157 (1817) (Fig. 3)
Description. Corm flattened-globose, 1320(30)
mm in diameter, tunics finely reticulate-fibrous, ex-
tending at the apex into a neck up to 60(100) mm
long. Cataphylls 35, white. Leaves (5)717, synan-
thous, occasionally hysteranthous but always devel-
oping immediately after flowers wither, 0.51.5 mm
wide, grey-green, glabrous or scabrid to papillose
along the margins and the keel beneath, with a white
stripe up to 1/3 of lamina width. Flowers 16, fra-
grant, pale pinkish-lilac to deep lilac-blue or purplish-
blue, usually darker-veined; throat white or lilac, more
or less pubescent. Prophyll present. Bract and bracte-
ole unequal, membranous, white, tapering at the apex
to long, rather flaccid tips. Perianth tube 4070(100)
mm long, white, lilac or purplish; segments (19)25
50 mm long, (5)816 mm wide, elliptic, oblanceolate
or obovate, acute or subacute, the inner segments of-
ten slightly smaller than the outer. Filaments 38 mm
long, white, glabrous to sparsely papillose-pubescent;
anthers 921 mm long, yellow. Style divided into three
red or occasionally orange branches, point of divi-
sion varying but usually found above the lower part
of anthers, each branch 315 mm long. Capsule ellip-
Fig. 3. Flowering plants
of Crocus pallasii subsp.
pallasii (a, b) and lat-
eral view of fower (c).
(Photographs by I. Syl-
lignakis and T. Karam-
plianis).
59 Phytol. Balcan. 19(1) Sofia 2013
soid, 1525 710 mm at maturity, carried at or just
above ground level. Seeds deep reddish to crimson,
subglobose to globose, 3.03.5(4) 2.53.0 mm. Ra-
phe usually a small ridge, wing-like, triangular, dis-
tinct on the side of chalaza, indistinct on the oppo-
site side. Caruncle small and irregularly formed, less
than 0.5 mm long and wide. Texture of seed smooth,
covered by velvet-like testa and a mat of long papillae.
Distribution and ecology. Crocus pallasii subsp. pal-
lasii is the most widespread member of the Cro-
cus series and extends from the F. Y. R. Macedonija
(Nikoli & al. 2010), Serbia (Ranelovi & al. 1990),
Bulgaria (Fgra & al. 2010; Assyov & Petrova 2012)
and Romania (Fgra & al. 2010) to the Crimea
(Seregin 2008) and Anatolia (Turkey), southwards to
Lebanon, Syria, Israel and South Jordan (Feinbrun
& Shmida 1977; Feinbrun-Dothan 1986; Mathew
1999). In Greece, C. pallasii subsp. pallasii was
known from the East Aegean Islands (Mathew 1982,
1983, 1984, 1988, 2000a; Christodoulakis 1986; Pan-
itsa 1997; Snogerup & al. 2001; Bazos 2005; Bazos &
Yannitsaros 2005) and from an unspecified record in
North Greece (Mathew 1999). We confirm the exist-
ence of this subspecies in the Greek mainland on the
basis of material collected from southeast Sterea El-
las to east Macedonia (see Appendix and Fig. 4).
Crocus pallasii subsp. pallasii grows in a variety of
habitats but mostly prefers calcareous grassland with
low vegetation, often amongst scattered Quercus coc-
cifera shrubs. It is also found in pine or oak woodland
openings, usually in shady places with fertile soil. It is
not a threatened subspecies in Greece, having been re-
corded from several localities with a good number of
individuals. As in the previous cases, however, an ex-
tensive collection of plants from the wild should be
strongly discouraged.
Taxonomic comments. Crocus pallasii subsp. pallasii
is a rather variable subspecies from a morphological
point of view. It is distinguished from its allies by the to-
tal style length which is up to half the length of perianth
segments, the concolorous (but often striated) perianth
that lacks any yellow band and the long corm neck that
reaches 60 mm or even 100 mm in Greek plants, de-
pending on the age of the corm. Most collections from
the mainland have extended necks,
usually 50100 mm long, and their
leaves are usually absent during
flowering. Their perianth segments
bear darker veins and measure 25
40 mm long and up to 14 mm wide,
longer and broader than the plants
from the East Aegean Islands. With
their long necks, the Greek plants
also resemble subsp. haussknechtii
(Boiss. & Reut. ex Maw) B. Mathew,
distributed in Northeast Iraq, West
Iran and Jordan. The leaves of the
southernmost mainland popula-
tion on Mt Imittos are synanthous
or begin to appear at the flowering
time, unlike the populations of Cen-
tral and North Greece and in com-
mon with the plants from the Aege-
an region.
Fig. 4. Distribution map of Crocus taxa in
Greece: Crocus biforus subsp. alexandri (green
circles); C. pallasii subsp. pallasii, known
localities (blue squares) and new localities
(red squares); C. reticulatus (brown triangles).
60 Karamplianis, T. & al. Floristic records and karyotypes of Greek Crocus
Mathew (2000a) commented on a Crocus popula-
tion of Samos Island, between Lazaros and Karvou-
nis summits of Mt Ambelos, and its similarity to C.
mathewii Kerndorff & Pasche, a species of Anatolia
(Kerndorff & Pasche 1994). We had the opportunity
to investigate this population in November 2009. The
Ambelos plants mostly have pale lilac segments and
some of them (but not all!) present the characteristic
violet throat of C. mathewii. However, on the basis of
morphological and karyological evidence, we do not
hesitate to assign the Samos population to C. pallasii
subsp. pallasii. The plants of Ambelos have well-de-
veloped leaves at flower ing time, measuring 0.7 mm
to 1.0 mm wide, slightly narrower than C. mathewii
(12 mm; Mathew 2000a). Their filaments are al-
so longer (58 mm vs. 34 mm long in C. mathewii)
and the corm tunics are evenly and finely fibrous-re-
ticulated along their whole length, whereas the tu-
nics of C. mathewii appear to have slender fibres ar-
ranged more or less in a parallel way at the lower part
of corm. These morphological differences between C.
pallasii and C. mathewii are by no means pronounced
and a clearer evaluation of the whole morphological
variation exhibited by the two taxa may be necessary
in the future. Karyological work revealed that the Sa-
mos population has a chromosome number of 2n =
14, which is different from 2n = 16 (Mathew 1999)
and 2n = 70 (Mathew 2000a, zhatay 2002) reported
for C. mathewii.
Together with Crocus asumaniae Mathew, C. pallasii
subsp. pallasii is apparently the most divergent member
of the Crocus series, a group that also comprises the cul-
tivated C. sativus (Grilli-Caiola & al. 2004). Recent phy-
logenetic studies (Petersen & al. 2008; Seberg & Peters-
en 2009) confirm its distance from the economically
important saffron crocus. Some of the important char-
acters that discriminate taxa within the Crocus series
are the relative style length, as compared to the length
of the perianth segments, the ratio of style length to
style branches, the perianth colour, the presence or lack
of indumentum on throat and filaments and the pres-
ence or absence of a neck at corm apex.
Karyology
Crocus reticulatus
Earlier works on the chromosomes of this species re-
port 2n = 10 (Mathew 1982), 2n = 12 (Brighton & al.
1973; Susnik & Lovka 1973; Baytop & al. 1975), 2n =
12 + 0 2B (Mathew 1982, Lovka 1995), 2n = 12 + 3B,
12 + 5B (Brighton & al. 1973), 2n = 14 (Popova 1972,
as Crocus variegatus Hoppe & Hornsch.; Mathew 1982;
Ranelovi & al. 2007), or 2n = 16 (Kuzmanov 1993,
as C. variegatus). The morphology of the karyotype is
seldom presented. We studied material from two lo-
calities of Mt Falakron (Table 1) and both showed a
diploid complement with 2n = 12 and a karyotype for-
mula of 2n = 2x = 2m + 10 sm (Fig. 5). No supernu-
merary chromosomes were found. The arm length ra-
tio of the chromosome pairs varies from 1.13 to 2.95.
The karyotype is classified as 3A, according to karyo-
Table 1. Chromosome data of the Greek Crocus species. Abbreviations: L/S = longest per shortest chromosome ratio; SA = Stebbinss
asymmetry index (Stebbins 1971); TF % = total form percentage index (Huziwara 1962); Ask % = ratio of long arm per total length
of chromosome set (Arano 1963); Syi % = mean length of short arms per mean length of long arms 100 (Greilhuber & Speta 1976).
A. Kar. = Andreas Karamplianis, Const. = Theophanis Constantinidis, Gorl. = Vassilis Gorlitsas, Tsif. = Spyros Tsiftsis, Th. Kar. =
Theophanis Karamplianis.
Taxon Voucher / Origin Location
coordinates
Alt.
(m)
Chromosome
number (2n)
Karyotype
formula
L/S SA TF
%
Ask
%
Syi
%
C. biflorus subsp.
alexandri
Th. Kar. & Tsif. 1841 /
Mt. Falakron
41 17.517' N
24 00.600' N
1210 8 6 sm + 2 st 2.17-3.29 3B 23.50-25.41 74.59
-76.50
30.73-34.07
C. pallasii subsp.
pallasii
Th. Kar. & A. Kar.
2050 / Mt. Imittos
37 57.465' N
23 49.055' N
900-920 14 6 m + 2 m-SAT +
6 sm
3.88-4.10 2B-2C 38.13-38.28 61.72-61.87 61.63-62.03
C. pallasii subsp.
pallasii
Th. Kar. & Const.1808
/ Samos Island
37 45.520' N
26 50.331' E
890-900 14 10 m + 4 sm 2.75-2.91 2B 36.07-37.83 62.17-63.03 56.45-60.86
C. pallasii subsp.
pallasii
Th. Kar. & Gorl. 1895
/ Mt. Vertiskos
40 46.620' N
23 15.857' N
575 16 2 m-SAT + 8 sm +
2 sm-SAT + 2 st +
2 st-SAT
3.12-3.52 2B-3B 22.85-34.39 65.61-77.15 29.41-41.80
C. reticulatus Th. Kar. & Tsif. 1839 /
Mt. Falakron
41 17.600' N
24 02.467' N
1470 12 2 m + 10 sm 1.83-1.88 3A 31.12-32.79 67.21-68.88 45.21-48.78
C. reticulatus Th. Kar. & Tsif. 1843 /
Mt. Falakron
41 17.517' N
24 00.600' N
1210 12 2 m + 10 sm 1.77-1.94 3A 31.85-31.97 68.03-68.15 46.75-46.98
61 Phytol. Balcan. 19(1) Sofia 2013
type asymmetry index (SA; Stebbins 1971). The long-
est per shortest chromosome ratio (L/S) ranges from
1.77 to 1.94. The total form percentage (TF %; Huzi-
wara 1962) varies from 31.12 to 32.79, the ratio of long
arm per total length of chromosome set (Ask %; Ara-
no 1963) ranges from 67.21 to 68.88 and the propor-
tion of mean length of short arms per mean length
of long arms (Syi %; Greilhuber & Speta 1976) varies
from 45.21 to 48.78 (Table 1).
The two subpopulations exhibit a very similar
chromosome complement.
Crocus biflorus subsp. alexandri
The chromosome numbers of 2n = 8 (Mathew 1982)
and 2n = 11 (Brighton & al. 1973) have been reported
for this subspecies; the latter was considered an aneu-
ploid karyotype from material of horticultural origin
(Alexandri). Our plants from Mt Falakron showed a
diploid complement with a rather asymmetrical kar-
yotype and a formula of 2n = 2x = 6 sm + 2 st = 8
(Table 1), lacking any supernumerary chromosomes
(Fig. 6). The relative length ratio of chromosome arms
varies from 1.92 to 3.72. The largest chromosome pair
is acrocentric (st), followed by three submetacentric
(sm) chromosome pairs, and the karyotype asymme-
try is classified as 3B (SA; Stebbins 1971). The longest
per shortest chromosome ratio (L/S) ranges from 2.17
to 3.29, the total form percentage (TF %) ranges from
23.50 to 25.41, the ratio of long arm per total length of
chromosome set (Ask %) ranges from 74.59 to 76.50,
and the symmetry index (Syi %) ranges from 30.73 to
34.07, as presented in Table 1.
Crocus pallasii subsp. pallasii
he chromosome numbers of 2n = 14 (Brighton & al.
1973; Brighton 1977; Randelovic & al. 2007; Candan
& al. 2009) and 2n = 16 (opova 1972) have been re-
ported for this subspecies. Both numbers were found
in Greek material but in different populations.
The plants from Macedonia (Table 1) have a com-
plement of 2n = 16 chromosomes (Fig. 4 a), classi-
fied as 2n = 2x = 2 m-SAT + 8 sm + 2 sm-SAT + 2 st
+ 2 st-SAT. Three pairs of satellites can be observed
on short chromosome arms and one more satellite on
the long arm of the longest chromosome (Fig. 7). The
arm length ratio varies from 1.30 to 4.18. The largest
chromosome pair is metacentric (m), with satellites
on both arms. The second in size chromosome pair
is acrocentric (st), with satellites on the short arms.
The remaining five chromosome pairs are submeta-
centric (sm), the largest one with satellites in the short
arms. The last in size chromosome pair is acrocen-
tric (st). The karyotype asymmetry (SA) is 2B or 3B
and the ratio of the longest per shortest chromosome
Fig. 5. Chromosome
metaphase plates of
Crocus reticulatus with
2n = 12 (a), scale bar =
10 m; karyotype (b),
scale bar = 10 m and
haploid idiogram (c),
scale bar = 1 m.
62 Karamplianis, T. & al. Floristic records and karyotypes of Greek Crocus
(L/S) varies from 3.12 to 3.52. The total form percent-
age (TF %; Huziwara 1962) varies from 22.85 to 34.39,
the ratio of long arm per total length of chromosome
set (Ask %) from 65.61 to 77.15 and the proportion of
mean length of short arms per mean length of long
arms (Syi %) from 29.41 to 41.80.
In contrast, material from the East Aegean Island
of Samos afforded 2n = 14 (Fig. 8), with a more sym-
metrical and quite different karyotype, as compared
to that of Macedonia. The karyotype formula is 2n =
10 m + 4 sm, without supernumerary chromosomes.
Relative length ratio of chromosome arms varies from
1.44 to 2.37. The largest chromosome pair is submeta-
centric (sm), followed by five metacentric (m) chro-
mosome pairs forming a karyotype asymmetry of 2B
(SA; Stebbins 1971). The longest per shortest chromo-
some ratio (L/S) varies from 2.75 to 2.91. Total form
percentage (TF %) varies from 36.07 to 37.83, the ra-
tio of long arm per total length of chromosome set
(Ask %) ranges from 62.17 to 63.03 and the symmetry
index (Syi %) varies from 56.45 to 60.86.
The population of Crocus pallasii subsp. pallasii
from Mt Imittos (Fig. 9) has the same chromosome
number with that of Samos. Its chromosomes are clas-
sified as 2n = 2x = 6 m + 2 m-SAT + 6 sm = 14. The rel-
ative length ratio of chromosome arms varies between
1.05 and 2.94 and the karyotype asymmetry (SA) is
2B or 2C. TF % is between 38.13 and 38.28, Ask % be-
Fig. 6. Chromosome
metaphase pl ate of
Crocus biforus subsp.
alexandri with 2n = 8
(a), scale bar = 10 m;
karyotype (b), scale
bar = 10 m and hap-
loid idiogram (c), scale
bar = 3 m.
Fig. 7. Chromosome
metaphase plate of Cro-
cus pallasii subsp. pal-
lasii from Macedonia
with 2n = 16 (a), scale
bar = 10 m; karyotype
(b), scale bar = 10 m
and haploid idiogram
(c), scale bar = 5 m.
63 Phytol. Balcan. 19(1) Sofia 2013
tween 61.72 and 61.87, and Syi % between 61.63 and
62.03 (see Table 1).
The differences in chromosome number and karyo-
type morphology between Greek populations of Crocus
pallasii subsp. pallasii are noteworthy. The population
of Macedonia has 2n = 16, thus confirming a report
by opova (1972) from the Central Balkan area. It al-
so exhibits a pronounced intra- and interchromosom-
al asymmetry (2B or 3B class in Stebbins classification,
high AsK value, low TF and Syi indices), as compared
to the 2n = 14 populations of Mt Imittos and Samos is-
land. The 2n = 16 karyotype cannot be explained by
simple chromosomal fission of a 2n = 14 complement,
because it also presents important chromatin reorgan-
ization which may be the result of shifts in centromere
position, additions or deletions and/or various translo-
cations. The existence of different cytotypes within C.
pallasii subsp. pallasii may be useful in circumscribing
groups within its natural distribution and offers cyto-
geographic markers that would shed light on the evolu-
tionary history of the species.
Acknowledgements. Many thanks to Erotokritos Kalogero-
poulos and Ioannis Syllignakis for offering photographs and data
on Crocus pallasii subsp. pallasii. Dr. Eleni Maloupa (National Ag-
ricultural Research Foundation, Thessaloniki), Dr. Nikos Krigas
(Aristotle University of Thessaloniki) and the staff of Ecology &
Systematics Department, Faculty of Biology, University of Athens,
provided facilities and support, which are gratefully acknowledged.
Fig. 8. Chromosome
metaphase plate of Cro-
cus pallasii subsp. pal-
lasii from Samos Island
with 2n = 14 (a), scale
bar = 10 m; karyotype
(b), scale bar = 10 m
and haploid idiogram
(c), scale bar = 1 m.
Fig. 9. Chromosome
metaphase pl ate of
Crocus pallasii subsp.
pallasii from Mt. Imit-
tos (Sterea Ellas) with
2n = 14 (a), scale bar =
10 m; karyotype (b),
scale bar = 10 m and
haploid idiogram (c),
scale bar = 5 m.
64 Karamplianis, T. & al. Floristic records and karyotypes of Greek Crocus
Crocus biflorus subsp. alexandri: Nomos Dramas,
Eparchia Dramas, Mt. Falakron, subalpine meadows
at the margins of Pinus forest, fertile substrate, marble,
with Crocus chrysanthus, Crocus reticulatus and Cro-
cus biflorus subsp. alexandri, 1470 m, 4117.600' N,
2402.467' N, 30.03.2010, Th. Karamplianis & S. Tsift-
sis 1840; ibid., at the margins and the openings of
Fagus and Pinus forest with Pulsatilla halleri subsp.
rhodopaea, fertile substrate, marble, 1210 m, Lat.
4117.517' N, Long. 2400.600' N, Th. Karamplianis
& S. Tsiftsis 1841.
Crocus pallasii subsp. pallasii: Nomos Dramas,
Eparchia Dramas, Korilovos hill, c. 1 km W of Kalli-
fitos village, clearings in Pinus brutia reforestations
and lowland grasslands with sparse Quercus coccifera
shrubs, limestone and alluvial deposits, c. 150200 m,
4110.630' N, 2411.890' N, 17.11.2012, S. Tsiftis (obs.
& photo); ibid., Mt Menikion, c. 1.3 km NW of Ka-
li Vrisi village, grasslands with sparse Quercus cocci-
fera shrubs, limestone, c. 300350 m, 4109.031' N,
2353.477' N, 10.11.2011 S. Tsiftis (obs. & photo). No-
mos Larissis, Eparchia Elassonos, c. 3.3 km NNW of
Elassona, along the road to Servia, margins of Pinus
woodland, close to the road, shady places with fer-
tile soil, c. 400420 m, 3954' N, 2210' N, 29.10.2007,
Th. Constantinidis & Th. Karamplianis 11619. Nomos
Samou, Eparchia Samou, Mt Ambelos, Samos Island,
c. 2.3 km NNE of Vourliotes village, c. 400500 m
NNE of Moni Vronta Monastery, small meadow in
clearings of Spartium junceum, Cistus sp. and decid-
uous shrubs, limestone, 500510 m, 3746.250' N,
2651.333' N, 20.11.2009, Th. Karamplianis & Th.
Constantinidis 1807; ibid., c. 7.5 km S of Vourliotes
village and c. 0.8 km E of Kioulafides settlement,
W of Lazaros summit, small meadows and mar-
gins of forest road in a burnt area with remnants
of Pinus nigra and Pteridium aquilinum, Crataegus
azorolus and Dittrichia viscose, limestone, c. 890
900 m, 3745.520' N, 2650.331' N, 20.11.2009, Th.
Karamplianis & Th. Constantinidis 1808; ibid., c.
1.5 km NW of Kioulafides settlement, close to the
road turn towards Tsouka, the eastern parts of Am-
belos summit, clearings of Pinus nigra forest, small
openings with Pteridium aquilinum, Cistus sp. and
Phlomis sp., mostly schist, with some limestone,
920930 m, 3746.109'N, 2649.271' N, 20.11.2009,
Th. Karamplianis & Th. Constantinidis 1809; ibid.,
c. 3.6 km S of Pirgos village, on the way to Spatha-
rei village, at roadsides in an earlier burnt area cov-
ered with Quercus pubescens, Rubus sp., Cistus cre-
ticus, Pyrus spinosa and grasses, schist, 580590 m,
3741.542' N, 2647.800' N, 21.11.2009, Th. Karam-
plianis & Th. Constantinidis 1820. Nomos Thessalo-
nikis, Eparchia Langada, Mt Vertiskos, c. 1 km before
Vertiskos town, meadows by the road, probably in de-
forested places of Quercus forest, perhaps psammitic
substrate, 680700 m, 4050.743' N, 2312.647' N,
02.11.2010, Th. Karamplianis & V. Gorlitsas 1839;
ibid., c. 2 km before Vertiskos town, meadows by
the road in a recreation place, probably deforested
places of Quercus forest and perhaps psammitic sub-
strate, 825 m, 4052.707' N, 2313.412' N, 02.11.2010.
Th. Karamplianis & V. Gorlitsas 1894; ibid., c. 2 km
after the junction to Areti village, on the road to-
wards the town of Sochos, meadows by the road,
semiruderal places, probably deforested places of
Quercus forest and perhaps psammitic substrate,
575 m, 4046.620' N, 2315.857' N, 02.11.2010,
Th. Karamplianis & V. Gorlitsas 1895. Nomos At-
tikis, Eparchia Attikis, Mt Imittos, c. 0.5 km be-
fore the first cluster of communication antennas at
the ridge of the mountain, fallow and waste places
by the road, limestone, 900920 m, 3757.465' N,
2349.055' N, 22.11.2012, Th. Karamplianis & A.
Karamplianis 2050.
Crocus reticulatus: Nomos Dramas, Eparchia Dramas,
Mt Falakron, 2.8 km NE of Xiropotamos village, mid-
altitude grasslands with Quercus coccifera, marble,
600800 m, 4112.802' , 2407.381' N, 26.02.2009,
S. Tsiftsis (obs. & photo); ibid., Mt Falakron, subal-
pine meadows at the margins of Pinus forest, fer-
tile substrate, marble, with Crocus chrysanthus,
Crocus reticulatus subsp. reticulatus and Crocus
biflorus subsp. alexandri, 1470 m, 4117.600' N,
2402.467' N, 30.03.2010, Th. Karamplianis & S.
Tsiftsis 1839; ibid., at the margins and the clear-
ings of Fagus and Pinus forest with Pulsatilla halleri
subsp. rhodopaea, fertile substrate, marble, 1210 m,
4117.517' N, 2400.600' N, 30.03.2010, Th. Karam-
plianis & S. Tsiftsis 1843.
Appendix
Plant material seen and populations used for cytological investigation (in bold).
65 Phytol. Balcan. 19(1) Sofia 2013
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