yes no Was this document useful for you?
   Thank you for your participation!

* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project

Document related concepts

Plant reproduction wikipedia , lookup

Glossary of plant morphology wikipedia , lookup

Flowering plant wikipedia , lookup

Ornamental bulbous plant wikipedia , lookup

Perovskia atriplicifolia wikipedia , lookup

Sarracenia wikipedia , lookup

Verbascum thapsus wikipedia , lookup

Crocus speciosus subsp. sakariensis
Jānis Rukšāns, Dr. biol. h.c.
June 2013
Published by the Alpine Garden Society © 2013
Abstract: Seven new taxa of Crocus L. from the Balkans and Turkey described.
Key words: Crocus speciosus subsp. sakariensis, Crocus speciosus subsp. bolensis,
Crocus speciosus subsp. hellenicus, Crocus speciosus subsp. elegans, Crocus vaclavii,
Crocus macedonicus, Crocus laevigatus subsp. pumilus.
Two of the species – C. speciosus and C. pulchellus – are placed by B. Mathew
in Series Speciosi. They occur around the Black Sea, in Turkey as far as its
Mediterranean coast, and in the Balkans. During this research we compared the
morphological features that allowed division of the C. speciosus complex into nine
different subspecies, four of them new. The crocuses studied are characterised
by parallel, fibrous, membranous corm tunics with more or less distinct basal
rings, leafless autumnal flowering (hysteranthous) and multi-branched stigmas.
Although some of the series proposed by B. Mathew were not clearly supported
by phylogenetic studies, Series Speciosi belongs to those which look very natural
and is well supported as a monophyletic group: it belongs to Section Nudiscapus
(Harpke, D. et al., 2012).
C. pulchellus occurs in W. Turkey, S. Bulgaria, the southern part of the former
Yugoslavia, N. Greece (Macedonia) and on some islands in the northern part
of the Aegean Sea – Thasos, Samotraki and Chios (Strid A., unpublished). It is
smaller in size than its closest relative C. speciosus and with a clearer, pale lilac-blue
background colour in the flowers. It was described in 1841 by the Rev. William
Herbert, who separated it from C. speciosus, with which it had been associated
by earlier authors. The most important morphological differences are a dark
yellow, even orange throat (somewhat similar are those of C. speciosus subsp.
xantholaimos, subsp. archibaldiorum, subsp. ibrahimii and subsp. sakariensis, but
of a paler shade), hairy filaments (papillose or nude in most forms of C. speciosus
except subsp. ibrahimii) and white anthers (usually yellow in C. speciosus, white in
subsp. ibrahimii and can be either in subsp. elegans). The flowers of C. pulchellus
have strong, delicate veining but they are never speckled or dotted on the outside
as is common in C. speciosus. C. speciosus has distinct secondary veining on the
perianth segments, not present in C. pulchellus. Generally C. pulchellus blooms
later than various forms of C. speciosus. It is a plant from moist meadows but it
grows in quite dry spots as well. Although being quite similar and requiring similar
conditions, they have never been recorded growing together in the wild (Mathew
B., 1982). Regardless of its wide range of distribution, all the collections of C.
pulchellus studied cytologically had 2n = 12 and the same karyotype (Brighton
C.A. et al., 1983).
The necessity to inspect the authentic samples from the locus classicus obliged
us to examine also the known populations in Asiatic Turkey. Samples from plants
grown and multiplied in cultivation for several generations are not completely
reliable because chance hybridization with other species growing nearby can occur,
especially when such closely related species as C. speciosus and C. pulchellus are
studied. In this study, plant material from 62 localities within the known area of
distribution of both species was compared, with the focus on C. speciosus as most
variable. The material from Armenia was provided by Armenian amateur gardener
Zhirair Basmajyan, who collected it in the wild. A few samples from S. Turkey had
been recently collected by Johan Nilsson from Gothenburg Botanical Garden.
In order to preserve the natural populations, from each locality of C. speciosus
sensu lato throughout its area of distribution, no more than 3-5 specimens were
collected for DNA research, herbarium and further observation in cultivation. To
examine the corm tunics, at several locations some 30-50 specimens were checked,
with exceptions when fewer specimens were available (Iran, Greece). After noting
the tunic features, the corms were left undisturbed and the soil was replaced,
avoiding damage to the populations. Our field research showed how important
it is to observe as many individuals as possible to be able to make an accurate
judgment about morphological features. We found that some features – such as
the structure of corm tunics (coriaceous or membranous), basal rings (distinct
or inconspicuous), tunic necks (long or very short, even absent), branching and
the position of stigma and the colour of the anthers – were very constant within a
taxon. Less constant was the throat colour. Some characteristics, such as the stigma
colour, were very variable – in the same population you could observe specimens
with yellow, orange and almost red stigmatic branches. In almost every population
2 JUNE 2013 Correspondence to: [email protected]
The genus Crocus consists of approximately 150 taxa (Harpke D. et al., 2012)
divided by B. Mathew (1982) into two subgenera (not supported by recent
phylogenetic research – Petersen G. et al., 2008; Harpke D. et al., 2012) and two
sections subdivided into 15 series. Later, one more series was added (Mathew B. et
al., 2009) and one series was moved to another section (Harpke D. et al., 2012).
The centre of distribution surrounds the Aegean Sea and most species are
concentrated in W. Turkey and the Balkans. Extensive research was conducted
in Turkey by Helmut Kerndorff and Erich Pasche and most of the new taxa
were described from just this region. It does not mean that crocuses from the
Balkans have been fully explored. Just recently a new species from Macedonia was
described, C. jablanicensis. Unfortunately, at present, we do not have good floras
for the region, but attempts are being made to rectify this. Arne Strid is working
on his Atlas of the Aegean Flora with distribution maps for all c. 3,600 species
of vascular plants, and I was greatly honoured to be invited to take part in the
mapping of Greek crocuses. During this research we found that several new crocus
taxa had not yet been described. These results are discussed in this publication.
examined, although the plants were very uniform we found 1-2 specimens that
stood out from the general description. This is quite common in the genus Crocus
and has been observed by H. Kerndorff et al. (2012) in the Crocus biflorus complex
as well.
Molecular studies allow us to identify crocuses with a high level of accuracy, but
this is of no use in the field. So we must rely on more substantial morphological
features to separate different crocus taxa. I used the features applied earlier by
Brian Mathew (1982) for distinguishing subspecies of C. speciosus – the structure
of corm tunics, the presence or absence of basal rings and elongated neck, the
degree of branching of the stigma and its position to the anthers, the colour of the
throat and anthers and the leaf dimensions. Each of the examined forms occurs
within a well-defined area and is more or less constant in its features.
Both C. pulchellus and C. speciosus are described as forming small cormlets
at the base of their bulbs. We found specimens with such cormlets in each wild
population (the only exception was the Iranian samples, where no cormlets were
observed in the wild, but we had access to a very limited number of specimens;
under cultivation cormlets were formed in abundance in sample WHIR-125, but
we suppose this to be an exception rather than a dominant feature). This trait
is prominently expressed in the old Dutch cultivars. A feature that we couldn’t
find mentioned in various earlier publications about C. speciosus but which was
observed in my collection of plants from Crimea, Iran (WHIR-125) and Turkey
(LST-374) was the formation of 7-10cm long lateral stolons with small cormlets at
the ends.
Another problem is determining where the border lies between the species
rank and the intraspecific ranks. This is not strongly defined by the Code. Many
botanists do not accept the subspecies rank at all. Brian Mathew in his monograph
(1982) lumped most of the crocuses with annulate tunics as subspecies of C.
biflorus. Recent molecular research by D. Harpke et al. (2012) does not support this
approach and the subspecies of C. biflorus sensu Mathew will be restored to species
rank, as well as the subspecies described later by H. Kerndorff and E. Pasche
(Kerndorff et al., 2012 and in preparation). The samples of C. speciosus examined
superficially were similar, so in my research I decided to keep them at the subspecific
level, but this can be changed when more data becomes available (DNA).
After analysing all the samples, I can separate nine well-defined areas with
distinct morphological features in each throughout the C. speciosus range. Having
put on the map the localities of the examined samples we can see that the type
subspecies is distributed generally in Crimea and the Caucasus, in the mountains
around the N. and E. coast of the Black Sea – Ukraine, Russia and Georgia. Some
specimens from Armenia (from Vahagni) do not quite fit within the general
concept of the type subspecies, having light yellow anthers and a bright yellow
throat. Although generally very similar morphologically, the Crimean plants
cytologically (2n=18) are different from the Georgian and Armenian samples
(2n=14). Plants from Armenia and Georgia have the widest leaves among all
C. speciosus sensu lato. The leaves in the samples from Crimea are narrower.
The Crimean plants have smaller chromosomes and a very different karyotype
(Brighton C.A. et al., 1983), therefore further research is required. At present we
cannot judge how far along the coast of the Black Sea the areas of the distribution
of the type subspecies extend. The N.E. part of Turkey was not included in this
research, but some samples from this area reported by Brighton et al. have a
different karyotype – 2n = 10.
I could not check C. polyanthus (nom. illeg.) described by A. Grossheim from
Azerbaijan Talish in 1936 and later brought under C. speciosus as a probable
synonym of the latter by Brian Mathew (1982). We (Rukšāns J., Seisums A.,
Zobova A.) did not find it in the vicinities of the village of Gosmeljan (from
where it was described) in May 1987 due to heavy overgrazing of the mountain
slopes there. Its description in the Flora Caucasica (Grossheim A., 1940) is too
incomplete but it could be conspecific with subsp. archibaldiorum described later
by me. According to A. Grossheim, the Talish samples have three stripes over the
backs of the petals – a similar pattern attracted my attention when I described
subsp. archibaldiorum. Subsp. archibaldiorum has a yellow throat and a shortly
branched stigma positioned between the anthers or overtopping them and has not
Distribution of Crocus speciosus s.l. and C. pulchellus in the wild. C. speciosus
(within red outline): 1 - subsp. speciosus; 2 - subsp. xantholaimos; 3 - subsp.
ilgazensis; 4 - subsp. archibaldiorum; 5 - subsp. ibrahimii; 6 - subsp. sakariensis;
7 - subsp. bolensis; 8 - subsp. hellenicus; 9 - subsp. elegans; 10 - C. pulchellus
(green outline). Blue outlines indicate areas where C. speciosus s.l. has been
reported but we had insufficient or no material for research. 11 - “C. polyanthus”.
the prolonged neck of the corm tunics. Five samples from Iran were researched by
Brighton C.A. et al. (1983) and they turned out to be cytologically different, having
2n = 12 and a distinctive karyotype, thus confirming my viewpoint in regarding
them as a different taxon.
From two passes in Sinop Province in N. Turkey comes subsp. xantholaimos. In
its corm tunic it is very close to the type subspecies. However, its throat in most
cases is bright yellow and the many-branched style is generally hidden among
the anthers. Another feature, confirming the identification, is the width of the
leaves, distinctly narrower than in the type subspecies. Our field observations at
locus classicus showed that its throat colour (by which it is named) is not always
distinctly yellow. Subsp. xantholaimos has 2n = 10, and the same number was
found in the samples from N.E. Turkey, too, but they were of a different karyotype
(Brighton C.A. et al., 1983).
Further west on the Ilgaz Dağ and Akdağ Mountains (near Amasya) it is replaced
by subsp. ilgazensis, which has a smaller flower than other forms and the fewbranched style is hidden among anthers. Compared with previously mentioned
subspecies it has very different corm tunics. They are thinly membranous, without
a neck, and the basal rings are very inconspicuous, so it is easily separable from
other subspecies. Plants from Ilgaz Dağ have 2n = 6. We did not examine Akdağ
near Amasya, although, according to Brighton C.A. et al., samples from there have
2n = 8, making them cytologically very similar, having only an additional small
submedian pair of chromosomes.
Further to the west, near Bolu and on the heights above Lake Abant, on almost
every mountain pass with more or less open clearings among trees and mountain
yailas, we found another crocus. This had coriaceous corm tunics with distinct
basal rings and a well-defined neck of old sheathing leaves, making it somewhat
similar to the type subspecies. But it was easy distinguishable from the latter by the
many-branched style hidden among the anthers. In this aspect it resembles subsp.
xantholaimos, but is separable by the throat colour, which is invariably pure white.
I decided to name it subsp. bolensis. It differs in the chromosome number, too.
Four samples researched by Brighton C.A. et al. from this part of the area had
2n = 8, separating this subspecies from other forms of C. speciosus.
Very distinct from all the other observed taxa is the form of C. speciosus
discovered by Ibrahim Sözen in the vicinities of Sakarya. In its corm tunic it is
close to subsp. ilgazensis but has a very bright yellow throat and a many-branched
stigma, which well exceeds the tips of the anthers. It is very unusual in the fact
that it grows at surprisingly low altitudes – 50m-120m (150m – by I. Sözen). Only
subsp. ibrahimii from Turkey in Europe is found at similar altitudes (100m-400m),
but it has white anthers. This complex of features allows us to regard this form as a
distinct subspecies of C. speciosus, which I decided to name subsp. sakariensis, after
the city closest to where it was found. Corms of this crocus lie in quite shallow
soil, only some 5-7(-10)cm deep, while the corms of other forms are positioned
 The table on the following pages gives an account of all observed populations
and their characteristic features, showing that all proposed subspecies are quite
easy to distinguish morphologically.
 With this knowledge in mind it was not very difficult to draw up a key for C.
speciosus subspecies and C. pulchellus (see page 10).
much deeper, to 20cm. Its flowers are comparatively small and in this aspect they
resemble those of subsp. ilgazensis.
Several travellers have seen C. speciosus in the extreme south of Turkey, almost
along its Mediterranean coast, from Bozkir in Konya province westward to
Altinyaka, west of Antalya. In autumn 2011 it was observed by a team from
Istanbul University, which did some research on its morphology (O. Erol et al.,
personal information). At present we have little information about the occurrence
of this form of C. speciosus, named here subsp. elegans for the pure white flower
tube seen in the first specimens, but the samples from Altinyaka colourwise more
resemble the typical C. speciosus forms, and similar ones were seen by J. Nilsson at
the locus classicus, too. Careful examination of the few specimens seen showed that
subsp. elegans has thin, membranous corm tunics with inconspicuous basal rings
and a long but very weak neck of old cataphylls. According to O. Erol, its anthers
are white, but this was not observed by J. Nilsson and me in the few samples seen:
probably the white colour was hidden by yellow pollen grains. Another possibility
is that the anthers observed by O. Erol were undeveloped. In such cases they can
be white-coloured. One sample from this district (S. of Beyşehir) was researched
cytologically by Brighton C.A. et al., who counted 2n = 18, the same as in the
Crimean samples but of a very different karyotype.
In 1983 Brian Mathew reported that C. speciosus had been observed in
Greece (Ipiros) but did not include this species in the taxonomic survey of
the Greek crocuses. According to A. Strid, C. speciosus was recorded in three
disjunct locations well separated from all other localities where C. speciosus has
been observed. Just recently I received a few samples of C. speciosus from the
southernmost population in Greece, collected by Jimmy and Karin Persson at
600m altitude, growing on north-facing mossy slopes. In autumn of 2012 at the
northernmost location in Greece, at an altitude of 1,300m just along the edges
of a yaila near trees, I observed another population of C. speciosus. Specimens
from both locations look very similar and by their floral characters they are not
separable from subsp. speciosus distributed in the Caucasus (locus classicus - “in
prov. Casp. Terek, Kour”). My field research showed that the Greek populations
of C. speciosus can be easily distinguished from the type just by the features of the
corm tunics. In the Greek samples they were thinly papyraceous, splitting parallely
at the base, with weakly developed basal rings and without a prolonged neck at the
apex. This allowed the Greek samples to be regarded as a distinct taxon, named
here as subsp. hellenicus.
Name /distribution /
altitude (observed)
of stigma
Stigma position
to anthers
Width of
1. subsp. speciosus
Crimea to Caucasus; 800-2,300m
up to
Crimea 18,
Caucasus 14
2. subsp. xantholaimos
N Turkey, Sinop prov.; 1,150-1,300m
below tips of
3. subsp. ilgazensis
N. Turkey, Ilgaz Dağ; 1,600-1,850m
with 6-8(10)
below tips of
membranous absent
6, 8
4. subsp. archibaldiorum
Iran; 650-2,000m
shortly, manybranched
at level of anthers coriaceous
or over-topping
5. subsp. ibrahimii
Turkey in Europe; 100-400m
short or
1.5-4(5)mm ?
6. subsp. nova sakariensis
N. Turkey, Sakarya; 50-150m
membranous absent
7. subsp. nova bolensis
N. Turkey, Abant, Bolu; 800-1,700m
below tips of
8. subsp. nova hellenicus
Greece; 600-1,300m
membranous present
but weak
9. subsp. nova elegans
S. Turkey, Akseki-Bozkir; 1,700m
membranous absent
but weak
Crocus pulchellus
N.W. Turkey, Turkey in Europe
& Balkans; 50-1,000m (1,800m –
Mathew B.)
few (6-9)
in tight
below tips of
* Width of leaves measured on cultivated flowering-size specimens, grown in same
conditions in author’s nursery.
JUNE 2013 9
Key for Crocus speciosus subspecies and C. pulchellus
1. Anthers yellow
2. Throat white or faintly yellow (in type subspecies)
3. Stigma many branched
4. Stigmatic branches well overtop anthers
5. Corm tunics coriaceous with distinct basal rings and long neck —
subsp. speciosus
5. Corm tunics membranous
6. Basal rings weak, tunic without elongated neck — subsp. hellenicus
6. Basal rings absent, tunic with long but weak neck — subsp. elegans
4. Stigmatic branches end at tip of anthers or below them — subsp. bolensis
3. Stigma with few branches hidden among anthers — subsp. ilgazensis
2. Throat yellow
7. Stigmatic branches end below tips of anthers, stigma with many branches —
subsp. xantholaimos
7. Stigmatic branches usually overtop anthers
8. Stigma with short branches ending at tips of anthers or overtopping them, corm
tunics coriaceous without neck — subsp. archibaldiorum
8. Stigma with long branches well overtopping anthers, corm tunics membranous
without neck — subsp. sakariensis
1. Anthers white
9. Stigma many-branched, well overtopping anthers
10. Corm tunics coriaceous, with basal rings — subsp. ibrahimii
10. Corm tunics membranous, without basal rings — subsp. elegans
9. Stigmatic branches 6-9, hidden among anthers — C. pulchellus
Crocus speciosus subsp. speciosus
Crocus speciosus subsp. speciosus
1. C. speciosus subsp. speciosus
Corm depressed – globose, 8-25mm in diameter, sometimes with cormlets at its
base, tunics coriaceous with distinct basal rings at the base and a long brown neck
of old cataphylls at the apex. Cataphylls 3-4, leaves 3-5, hysteranthous, emerging
long after flowering, glabrous or with a ciliate or scabrid margin or pubescent on
the upper lamina, 4-8(10)mm wide and up to 37cm long, without ridges in lateral
channels. Flowers 1-2, fragrant, lilac blue usually conspicuously veined darker,
usually with distinct secondary veining, sometimes silvery or whitish on the
outside or with a darker speckling. Throat glabrous, whitish or faintly yellow, rarely
yellow (Armenia). Prophyll absent, bract and bracteole subequal, membranous.
Crocus speciosus subsp. speciosus
An albino form of Crocus speciosus
subsp. speciosus (Crimea)
10 JUNE 2013 Crocus speciosus M. Bieb., Beschr. Land. Terek Casp. 129 (1800)
Habitat of Crocus speciosus subsp. xantholaimos at Sinop in N. Turkey
Flowers and corm of Crocus speciosus subsp. xantholaimos
Perianth tube 5-20cm long, white or purplish toned in upper part, segments 3-7cm
long, inner distinctly broader than outer up to 2.2cm wide. Filaments white or
light yellow, glabrous or minutely papillose, 4-11mm long, anthers yellow, 2.5-3
times longer than filaments. Style divided at the top of anthers into many yellow
to deep orange slender branches well overtopping anthers. Capsule carried just
above ground level at maturity, seeds reddish-brown, nearly globose, up to 3mm in
diameter. 2n = 14 (Caucasus), 18 (Crimea).
Habitat: Woods or alpine meadows, clearings in forest on limestone or noncalcareous formations, altitude: 800m-2,500m.
Flowering period: September to November.
Type locality: Caucasus mountains (“in prov. Casp. Terek, Kour”).
Distribution: Crimea (Ukraine) and Caucasus (Russia, Armenia, Georgia,
Azerbaijan?, Turkey?)
2. C. speciosus subsp. xantholaimos Mathew, The Crocus: 111 (1982).
The general description is the same as that of subsp. speciosus but with the
following characteristics: leaves narrow (1-2.5mm wide), throat of perianth yellow,
sparsely pubescent, style much branched but shorter than anthers. 2n = 10
Habitat: Clearings in Abies and Rhododendron forest, alpine turf (yaila) and stony
hillsides on limestone formations, altitude: 1,150m-1,300m.
Flowering period: September to October.
Flower and corm of Crocus speciosus subsp. ilgazensis
Type locality: Turkey, Sinop prov., passes between Kabali and Boyabat.
Distribution: N. Turkey (Sinop).
3. C. speciosus subsp. ilgazensis Mathew, The Crocus: 111 (1982).
The general description is the same as that of subsp. speciosus but with the
following characteristics: corm tunic membranous, lacking distinct basal rings.
JUNE 2013 13
Flowers generally smaller, tube up to 5-8cm long, segments up to 3.5cm long. Style
divided in 6-8 branches hidden among anthers. Seeds c. 1.5mm diameter.
2n = 6, 8.
Habitat: Clearings in Abies forest, turf on stony hillsides on limestone formations,
altitude: 1,600m-1,850m
Flowering period: September to October.
Type locality: Turkey, Çankiri prov., Ilgaz Dağ pass, 1,700m.
Distribution: N. Turkey (Çankiri and Amasya).
4. C. speciosus subsp. archibaldiorum Rukšāns, The Alpine Gardener: 80: 209
The general description is the same as that of subsp. speciosus but with the
following characteristics: flowers whitish from the outside with minutely branched,
comparatively wide lilac stripes from the bottom of the segments up to the tip,
the perianth throat yellow, stigmatic branches short, ending at the level of or
overtopping the anthers, leaves develop soon after flowering, corm tunics without
a prolonged neck. 2n = 12.
Habitat: Steep mountain slopes in turf or at Fagus forest sides and among shrubs,
on limestone formations, altitude: 650m-2,100m
Flowering period: October.
Type locality: Iran, Kuhha-ye Tales, between Nav and Khalkhal, steep mountain
slopes just before pass, 2,080m.
Distribution: N.E. Iran (Mazandaran, Gilan,), S. Azerbaijan? (as C. polyanthus
Grossheim, nom. illeg.).
Flowers and corms of Crocus speciosus subsp. ibrahimii
5. C. speciosus subsp. ibrahimii Rukšāns, The Alpine Gardener: 80: 210 (2012).
The general description is the same as that of subsp. speciosus but with the
following characteristics: perianth throat yellow, filaments hairy, anthers white.
From the similar C. pulchellus it is separable by the position of the stigma (well
overtopping the anthers), form and pattern design of the tepals. 2n=?.
Habitat: Clearings in Quercus forests and heath on limestone or non-calcareous
formations, altitude: 100m-400m.
Flowering period: October to November.
Type locality: Turkey in Europe, near Çanakça
Distribution: Turkey in Europe (Yildiz Dağ). Perhaps also found in adjacent S.E.
6. C. speciosus subsp. sakariensis Rukšāns, subsp. nov., a subsp. specioso fauce
lutea, cormi tunico papyraceo, basi in fibras parallelas fisso sine collum, floribus
minoribus differt. A subsp. ilgazensis et xantholaimos stylus ad apices antherarum
divisus ramosissimus differt. Typus: Turkey, S.E. of Sakarya, foothills of Elmacik
Dağ, 2012-10-31, alt. 85m. Holo: Gatersleben, GAT 19548; Iso: GB.
Habitat and corm of Crocus speciosus subsp. sakariensis (see front cover)
Habitat of Crocus speciosus subsp. bolensis in Turkey
Flower and corm of Crocus speciosus subsp. bolensis
The general description is the same as that of subsp. speciosus but with the
following characteristics: corm tunics membranous, without basal rings, perianth
tube up to 10cm long, perianth segments 30-35mm long, throat deep yellow.
Stigma many-branched, well overtopping anthers. Leaves 3-6mm wide. 2n=?.
Habitat: In shade under Corylus and among young plantings in full sun, on
humus-rich gravelly clay at 50m-150m altitude.
Flowering period: October to November.
Type locality: Turkey: Sakarya
Distribution: Known only from the type locality.
Habitat: On open alpine turf (yaila) and in clearings of Pinus and Abies forests,
altitude: 850m-1,700m.
Flowering period: October to November.
Type locality: Turkey: Heights above Lake Abant
Distribution: Turkey: Abant, Bolu and Gokceler Dağ.
7. C. speciosus subsp. bolensis Rukšāns, subsp. nov., a subsp. specioso stylus ramus
inferus apices antherarum finis differt, a subsp. ilgazensis stylus ramosissimus, a
subsp. xantholaimos faucis albus differt. Typus: Turkey, Bolu, heights above Lake
Abant. Holo: Gatersleben, GAT 19558.
The general description is the same as that of subsp. speciosus but with the
following characteristics: stigma well-branched but ends below the tips of anthers.
Flowers are smaller than in the type subspecies, generally nicely striped. From
subsp. ilgazensis well separable by the distinctly many-branched stigma, from
subsp. xantholaimos by the white throat. 2n=8.
8. C. speciosus subsp. hellenicus Rukšāns, subsp. nov., a subsp. specioso cormi
tunico papyraceo basi in fibres parallelas fisso, cum tunica basalis indistinctus
(hebdomadalis evolutus), sine collum differt. Typus: Greece, Ioannina, Vikos
Canyon, nr. Monodendri. 2012-10-12. Holo: Gatersleben. GAT 19551.
Fokida: GAT 20316 (Greece, nr. Varnakovo Monastery, ex culturae in horto Jānis
Rukšāns, 2012-11-07).
The general description is the same as that of subsp. speciosus but with the
following characteristics: corm tunics membranous, with weakly developed basal
rings, without prolonged neck. Leaves narrow, 1-3(4)mm wide. 2n=?.
Habitat: On yailas at forest edges and on mossy slopes, on limestone; altitude:
Flowering period: October to November.
Type locality: Greece, Ioannina, Vikos Canyon, nr. Monodendri.
Two forms of Crocus speciosus subsp. hellenicus from different locations
Three images of Crocus speciosus subsp. elegans and, below, its habitat
Distribution: Greece. Three disjunct localities in Ioannina, Etoloakirnania and
9. C. speciosus subsp. elegans Rukšāns, subsp. nov. a subsp. specioso antherae
albae(?) et cormi tunico papyraceo basi in fibres parallelas fisso, cum tunica
basalis indistinctus sed cum collum papyraceo differt. Typus: Turkey, Konya prov.,
Esereyrek Da. Holo: GB
The general description is the same as that of subsp. speciosus but with the
following characteristics: corm tunics membranous, splitting longitudinally into
stripes, basal rings indistinct. Long neck formed by old cataphylls present, but thin
and very brittle, so careful digging is needed for observation. Anthers white (?).
Leaves up to 4(?) mm wide. 2n = 18.
Habitat: On clearings and edges of Abies forests and rocky outcrops, altitude:
Flowering period: October to November.
Type locality: Turkey: Konya province, south of Suğla Gölü.
Distribution: Turkey: Geyik Dağlari (known from two localities) and Görece
Daği(?) and possibly westermost end of Akçali Dağlari (KPPZ-9022B).
JUNE 2013 19
Arne Strid supplied me with information about all gatherings of crocuses known
to him from Greece. This allowed me to make further research about some
crocuses with an unclear position at present. This research is not finished yet and
more fieldwork is necessary in order to obtain material of some new or possibly
forgotten species. One such is Crocus athous Bornm. described in 1944 but later
included under C. sublimis Herb. It was rediscovered by A. Strid at its locus
classicus (personal information) and, according to the material available at present,
seems to be a valid species, but still some research is needed to confirm its status.
In the same part of Greece was photographed (under the name of C. pallasii)
a very interesting crocus gathered earlier (1914, 1944) but misidentified as C.
alexandri (C. biflorus subsp. alexandri) or C. biflorus. In the pictures seen it looked
very different from C. alexandri as well as from C. stridii, which was growing in
the vicinity. But identification of crocuses by pictures especially in the C. biflorus
group is almost impossible, so a new gathering of this crocus was necessary to
decide on its status. The first attempt to find it in February 2012 failed due to a
heavy storm at sea, which prevented me from reaching Athos. In early March 2013,
with my friend Vaclav Jošt (Czech Republic), I revisited the possible localities of
this crocus and, after a three-day search, found a small population. We gathered a
few samples for a cytological investigation at the Leibniz Institute of Plant Genetics
and Crop Plant Research in Gatersleben. I decided to name it Crocus vaclavii after
Vaclav Jošt, a leading lily breeder in the Czech Republic, in recognition of his help
in discovering it.
The corm of C. vaclavii is globose, around 10-15mm in diameter. Tunics are
coriaceous, old tunics very hard, the new ones thinner, greyish brown with
prominent basal rings. The edge of the rings is uneven but without prominent
teeth. The tunic neck is short, not longer than 3mm. Cataphylls 3-4, prominent,
silvery white, rarely slightly greenish shaded. Leaves much longer than flowers at
anthesis, usually 3 or 4, rarely 2 or 5, on average 3.5 (from 45 samples recorded),
dark green (2)2.5-3(3.5)mm wide with a prominent white stripe around one
third of the leaf diameter or slightly less, without ribs underneath. The bract and
bracteole are present, subequal. The perianth tube is white, in the upper part with
greyish-purple stripes. The throat is nude, yellow with a greyish flush at the edge,
more prominent in the middle of the petals, giving a somewhat muddy impression.
The perianth segments on the inside are invariably bluish-violet. The outer
segments are 24-33mm long (usually 29-30mm) and 10-14mm (usually 13mm)
wide, mostly with 3 prominent dark purplish violet stripes along the length of the
petals, those at the side usually feathered. Stripes rarely reach only one-third of the
petals’ length or the outside is speckled. We observed one specimen with yellowshaded segment backs. The inner segments are smaller – up to 3mm shorter and
1mm wider than the outer ones, bluish violet with a narrow dark grey “tongue” at
the base, bordered in yellow on the sides, but with a diffused white zone at the top.
Filaments are yellow, 8-10mm long, distinctly papillose, anthers 11-15mm long,
The habitat of Crocus vaclavii at Athos and, below, two images of the species
JUNE 2013 21
yellow or yellow with a black connective (in around 20% of observed specimens).
The style is yellow, gradually becoming orange to orange-red at the top, usually
divided into three short (up to 5mm long) papillose branches ending at the tip of
the anthers or slightly overtopping them. Rarely the branches are shorter and end
below the anthers (observed in 4 specimens of the 45 measured). The capsule and
seeds were not seen. The chromosome number is unknown.
Type specimen: this was collected on a very wet seaside meadow on the east
coast of the Athos Peninsula at only 5-10m above sea level, where it was growing
in coarse gravelly soil formed from decomposed granite and probably received
salt-water spray during heavy sea storms. It was cited by Bornmüller in 1944, most
likely from the same locality, as Crocus biflorus. The same species was observed
under cultivated Corylus plantings at 650m near Karyes in 1914 (as C. biflorus
alexandri) and it was pictured near Iviron Monastery in 2007 (as C. pallasii –
published on the website of Naturetrek).
The leaf morphology is an important feature in distinguishing crocus species.
This new taxa belongs to the species in the “biflorus” group without ribs in the
grooves on the underside of the leaves. In this it resembles C. stridii, C. biflorus
(type), C. alexandri and the recently described C. bifloriformis and C. babadagensis.
In the black colour of its anthers the new species mostly resembles C. stridii, which
grows in similar conditions in the same region of Greece (Chalchidiki) although
both are well separated geographically, but C. stridii is much leafier, having 5-8
leaves, and its throat is papillose. All specimens of C. stridii seen by me had flowers
with a white base colour. The flowers of C. alexandri invariably have white throats.
More difficult is to separate it from other species without ribs in the leaf grooves.
C. biflorus sensu stricto has narrower leaves (only 0.5-2.0mm wide), its anthers are
shorter (5-11mm long) and it grows on limestone-based formations. Compared
with C. bifloriformis and C. babadagensis from the opposite coast of the sea (W.
Turkey), the new species is less leafy and its flowers are blue coloured on the
outside and within.
Crocus macedonicus
Flowering period: January to March.
Type locality: Greece, Macedonia, E. coast of Athos Peninsula.
Distribution: Greece, Athos Peninsula. A very local endemic.
Crocus vaclavii Rukšāns species nova. Cormi tunicae coriacea cum annulis ad
basem. Annuli sine manifeste dentes. Folia 2-3, 5-5 (n = 45), 2.5-3mm lata, glabra,
subter sine costis, bene evoluta ad florationem. Corolla fauce aurantiaca, glabra.
Segmenta caerulea. Segmenta exteriora extus plerumque striata vel pinata violaceo,
raro maculata. Filamenta saturate lutea, 8-10mm longa, papillosa. Antherae 1115mm longae, luteae vel cum connectivo nigro vel griseo (20%). Stigma antheris
plerumque aequalis, rami stigmatici 5mm longa. Capsula non visa. Typus: Greece,
Macedonia, Athos, seaside wet meadow at only 5-10m above sea level, coarse
gravelly soil formed from decomposed granite, 03-03-2013. Holo: GAT 23017. Iso:
Habitat: on decomposed granite-based seaside meadows and higher in grass;
altitude 5m-650m.
In 2002 David Stephens published a report about crocuses seen in central and
eastern Macedonia in Greece (Crocus Group Bulletin, No. 30). His team followed
in the footsteps of Neil Jacobsen, who had reported seeing a small population of
C. pallasii between Drama and Xanthi and between Serres and Thessaloniki. Later
the same crocus was reported by Christopher Greenwell and Simon Silcock (2010)
on the internet (Scottish Rock Garden Club Forum). In February 2012, with my
son-in-law, I visited Greek Macedonia in search of the already described Crocus
vaclavii. Because of a heavy storm, we could not reach our destination and used
the time to explore the continental part of Greece. Near the village of Ossa, among
flowering C. chrysanthus, we discovered an autumn-blooming crocus (by leaves)
which turned out to be conspecific with the above mentioned crocus, which at first
glance resembled C. pallasii but differed in several important features.
Brian Mathew recognises four subspecies (at present regarded as separate species,
Erol O. et al., in preparation) of Crocus pallasii, and in order to separate them he
uses the shape of the flower segments and the features of the corm tunics. Three
of them are distributed in S.E. Turkey and adjacent countries. The greatest area is
occupied by the type subspecies which occurs in Crimea (type locality), Bulgaria,
Romania, Serbia, on Greek Islands (Chios, Samos) and in W., Central and S.
Turkey. By its locality the Macedonian crocus could belong to the type subspecies,
which is characterised by a short neck formed by old cataphylls (up to 2cm long),
but the plants checked by us had a much longer and very distinctly fibre-like neck.
In examined samples (35) its neck length varied from 5-10.5cm but mostly
averaged 6-8cm. So a long neck is seen only in subspecies distributed very far to
the east (subsp. turcicus has 3.5-6cm; subsp. haussknechtii up to 10cm long, and
subsp. dispathaceus 3-7cm long). Further research showed that this crocus was
less leafy than C. pallasii. All acquisitions of C. pallasii sensu lato in my collection
(39 samples) have 10-17 leaves per shoot, while the plants from Macedonia have
only 5-6 leaves per shoot. In spring 2013 we found a large population on an
abandoned, earlier cultivated field with deep rich soil. The plants here were leafier,
with up to 9 leaves with an average number of 6.5 (from 40 studied specimens)
and the highest number of seed pods (flowers) per plant was 3. The few collected
corms bloomed in my collection very late, in December, when all C. pallasii
sensu lato had long finished, but this probably was caused by the early collecting
in the wild and the disturbance of the normal course of development. In flower
they were indistinguishable from C. pallasii. Preliminary results of the molecular
investigation (unfinished) by Dörte Harpke from Gatersleben Institute confirmed
that this crocus was different from C. pallasii. So there is no doubt that it is a new,
undescribed species which I have decided to name C. macedonicus.
Crocus laevigatus subsp. pumilus from Vouvala, Crete
Crocus macedonicus Rukšāns species nova. Cormi tunicae fibrosis apice cum
collum (5-)6-8(-10.5) centrimetrum longum plerumque persistens. Folia (4)57(-9), synantha, 1(-1.5)mm lata, margine glabra. Flores autumnales, plerumque
solitarii (-3) Perigonium saturate lilacino-coeruleum, prope basin valde purpureo
venosum. Fauce album. Filamenta alba, antherae flavae. Stylus in ramos tres
stigmaticos, rubros, apice expansos divisus, quam antherae aequans or superans.
Crocus pallasii affinitas sed collum longum et folia numero minor bone differt.
Typus: Greece, Macedonia, N.E. Ossa. Holo: GAT 20314 (ex culturae in horto Jānis
Rukšāns, 2012-12-08); Iso: GAT 20315.
Habitat: on limestone-based mountain slopes in short turf; altitude 350m-700m.
Flowering period: October to November.
Type locality: Greece, Macedonia, N.E. of Ossa
Distribution: Greece, Macedonia, Vertisko ridge. The latest research shows that it
is more widespread than had been supposed earlier. We found it at every stop on
the southern slopes of Vertisko ridge in a westerly direction, starting from Sochos.
Locally it was very abundant: on abandoned cultivated field its leaves covered the
ground like grass.
While working on my collection of Greek crocuses, my attention was focused
upon C. laevigatus represented by 28 gatherings from various localities. Several
pots (8 gatherings) looked distinctly smaller in flower than others. Having checked
the labels I found that they were all from Crete, while other samples were from
the mainland or other islands. Then I remembered a note in Brian Mathew’s
monograph (1982) about diminutive, odourless forms of C. laevigatus on the
island of Crete, stating that an “overlap in measurements is too great to be of value
taxonomically”. Those observations were made of wild growing specimens or of
herbarium specimens of wild plants. The cultivation of plants in a more or less
identical situation (i.e. in culture) allows us to compare properly the measurements
of various morphological features, for in the wild they are subject to the pressures
of natural conditions that vary from season to season and depend on the vagaries
of the weather. This differs in each place and does not always allow the genetic
potential to express itself fully.
While checking the flower measurements on all my gatherings of C. laevigatus
grown in identical conditions, with the same substrate, watering and fertilizing
regime, I found that flower dimensions were significantly different. The petal
lengths in cultivated Cretan plants varied from 20-26mm, but in the gatherings
from other localities they were 30-41mm long (Brian Mathew states that they vary
from 1.3-3cm in length, so under cultivation flowers become bigger). Furthermore,
all the Cretan plants were odourless while others were wonderfully fragrant. It
means that the Cretan plants can be regarded as sufficiently different to give them a
subspecies status.
Crocus laevigatus subsp. pumilus Rukšāns subsp. nova a subsp. typica flores
minoribus et inodori bone differt. Typus: Greece, Crete, Omalos Plateau, alt. 940m,
ex culturae in horto Jānis Rukšāns, 2012-10-23. Holo: GAT 20317. Iso: GB
Habitat: open stony and rocky places or in sparse scrub; altitude 10m-1,520m
Flowering period: October to December.
Type locality: Greece, Crete, Omalos Plateau.
Distribution: Greece: the island of Crete – widespread throughout the island.
I thank Dörte Harpke for her assistance in cytological research, the Turkish
amateur botanist Ibrahim Sözen for sharing with me his knowledge about Turkish
crocuses, Arne Strid and David Stephens for the information about Greek crocus
localities, Osman Erol for data about subsp. elegans, Johan Nilson and Henrik
Zetterlund for living material and pictures (J. Nilson – subsp. elegans in wild) of
Greek (subsp. hellenicus) and Turkish crocuses (subsp. elegans), and especially
Christopher Greenwell and Simon Silcock for sharing their observations about
C. macedonicus. To Dmitriy Zubov (Ukraine) I express my thanks for assistance
during the field work in Crimea, to Zhirair Basmajyan (Armenia) for the living
material from Armenia, and to Vaclav Jošt (Czech Republic) and Krišs Karnītis
(Latvia) for assistance in discovering the new crocus on the Athos Peninsula and
Macedonia (Greece).
Brighton, C. A., Mathew, B., Rudall P. 1982. A Detailed Study of Crocus speciosus
and its ally C. pulchellus (Iridaceae). Pl. Syst. Evol. 142: 187-206 (1983).
Davis, P. H., ed. 1984. Flora of Turkey and the East Aegean Islands. Vol. 8.
Edinburgh University Press.
Grossheim, A. A. 1940. Flora Caucasica. Vol. 2, 2d ed. Baku: AzFAN. (In Russian)
Harpke, D., et al. 2012. Phylogeny of Crocus (Iridaceae) based on one chloroplast
and two nuclear loci: Ancient hybridization and chromosome number evolution.
Mol. Phylogenet. Evol.
Kerndorf H., Pasche E., Harpke D. & Blattner F. 2012. Seven New Species of
Crocus (Liliiflorae. Iridaceae) from Turkey. Stapfia 97: 3-16.
Mathew, B. 1982. The Crocus: A Revision of the Genus Crocus (Iridaceae). London:
B.T. Batsford.
Mathew, B. 1983. The Greek Species of Crocus (Iridaceae): a taxonomic survey.
Crocus laevigatus subsp. pumilus from the Omalos Plateau, Crete
Ann. Musei Goulandris. Vol. 6: 63-86.
Mathew, B., Pettersen, G., Seberg, O. 2009. A reassessment of Crocus based on
molecular analysis. The Plantsman, New Series vol. 8: 50-57.
Maw, G. 1886. A Monograph of the Genus Crocus. London: Dulau.
Petersen, G. et al. 2008. A phylogeny of the Genus Crocus (Iridaceae) based on
sequence data from five plastid regions. Taxon vol. 57: 487-499.
Rukšāns, J. 2010. Crocuses: A complete guide to the genus. Portland, London:
Timber Press.
Rukšāns, J. 2012. A revision of Crocus speciosus in Turkey and Iran. The Alpine
Gardener vol. 80 (2): 206-211.
JUNE 2013 27
Additional copies of this pamphlet may be obtained from the
Alpine Garden Society, the international organisation devoted
to the cultivation, conservation and exploration of alpine and
rock garden plants. To place an order, visit our website at the
address below and follow the link for the Book Shop.
You can also become a member of the Society through our
website. Recruiting new members is vital to our work and
enables us to produce pamphlets and other publications such
as this one.
Crocus vaclavii
Crocus macedonicus
Our members enjoy many benefits including:
 The Alpine Gardener, our highly regarded 132-page quarterly
 Access to our annual Seed Exchange, with around 5,500
different seeds listed.
 Botanical tours in many different parts of the world, led by
expert guides.
 A wide range of discounted books including field guides,
monographs and practical volumes, some of them difficult to
obtain elsewhere.
 In the UK and Ireland, we organise 23 National Shows, some
featuring more than 800 plants grown by our members.
 AGS News, our quarterly newsletter, gives information on our
shows, plant fairs, conferences and other events.
 Our website contains the AGS Encyclopaedia of Alpines, an
unrivalled reference work on the mountain plants of the world.
For plants people
Crocus speciosus subsp. speciosus
Crocus speciosus subsp. bolensis