Phytotaxa 10: 17–25 (2010) ISSN 1179-3155 (print edition)

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Copyright © 2010 • Magnolia Press ISSN 1179-3163 (online edition)

Tulipa albanica (Liliaceae), a new species from northeastern Albania

LULËZIM SHUKA1, KIT TAN2 & SONJA SILJAK-YAKOVLEV3

1
Department of Biology, Faculty of Natural Sciences, Tirana University, Bld. ZOG I, Albania
2
Institute of Biology, University of Copenhagen, Øster Farimagsgade 2D, DK-1353 Copenhagen K, Denmark. E-mail: kitt@bio.ku.dk
3
Univ. Paris-Sud, Lab. Ecologie, Systématique & Evolution, CNRS UMR 8079, AgroParisTech, Bât. 360, 91405 Orsay, France

Abstract

Tulipa albanica (Liliaceae) is illustrated and described as a new species from a serpentine area in Kukësi district,
northeastern Albania. It is compared with T. scardica from the western Balkans (Macedonia and Kosovo) and with T.
schrenkii from the Ukraine, Caucasus, Central and Southwest Asia. The chromosome number, karyotype features and
nuclear DNA content (2C-value) were assessed by karyological and flow cytometry techniques and provided for the first
time for this species. The chromosome formula is 2n = 2x = 24 (2 metacentric, 3 submetacentric and 7 subtelocentric
chromosome pairs) and the genome size 54.15 (± 0.23) pg. Two different morphotypes of T. australis were noted as
occurring on limestone or serpentine in Albania.

Key words: Balkan Peninsula, chromosome number, nuclear genome size, serpentine soil, Tulipa australis,
morphotypes

Introduction

Tulipa Linnaeus (1753: 305) is a large genus with ca. 100 (Hall 1940) or 113 species as recently accepted by
Govaerts (2010). The species are widely distributed from northeastern China and Japan to Central and
Southwest Asia, North Africa and Europe (Botschantzeva 1982, Bermejo & Sánchez 2009). The triangle
between the Tien Shan and Pamir-Alai mountain ranges in Central Asia is considered to be the main centre of
diversity and the Caucasus region a secondary centre (Terzioğlu & Coşkunçelebi 2002, Zonneveld 2009). The
Iranian Plateau is home to at least 36 species of wild tulips (Ghahreman et al. 2007). The number of native
tulip species found in the Balkan Peninsula is much less, varying from c. 15 (Hayek 1933) to 22 (Govaerts
2010). Eighteen species have been recorded from European Turkey and Anatolia (Terzioğlu & Coşkunçelebi
2002, Coşkunçelebi et al. 2008, Eker & Babaç 2010).
Except for the cultivated tulips of garden origin, the genus is in Albania represented by only one native
species: T. australis Link (Schrader 1799: 317) that was previously known as T. grisebachiana Pantocsek
(1873: 265) and was first collected on the limestone mountain Mt Tomorri by Markgraf in 1928 (Markgraf
1931). Tulipa grisebachiana was originally described from Mt Gliva (near Trebinje, Hercegovina). Tulipa
australis was also reported as T. sylvestris subsp. grisebachiana and subsp. celsiana by Hayek (1932–1933:
71, syn.: T. celsiana DC. in Redouté 1803: t. 38). It is not apparent whether Hayek was referring to plants
from limestone or from serpentine substrate. Photographs of T. australis from both serpentine (e.g., Surroj
area, Fig. 1A) and limestone substrate (e.g., Mt Çajupi, NE of Gjirokaster, Fig. 1B) are here provided. They
represent two different morphotypes with slight differences in perianth shape, size and colour. We have
maintained T. grisebachiana and T. celsiana in synonymy of T. australis and have not formally recognized the
two morphotypes from differing substrates. We here report the discovery of a second and undescribed tulip
from Albania belonging to subg. Tulipa sect. Tulipa; this tulip has both red and yellow flowering forms.

Accepted by M. Christenhusz: 5 Oct. 2010; published: 28 Oct. 2010 17

 During a visit on 22 May 2008 to the meadows near Surroj village in Kukësi district in northeastern
Albania, two tulip plants with strongly undulate leaves were observed by L. Shuka. They were bearing young
fruits and no flowering plants were seen. He asked Luan Vata, a fourth year biology student at Tirana
University living in Surroj village, who was investigating Crocus in the area, to look out for flowering
specimens of this tulip. In early April 2009 Vata reported that the beautiful tulip had been found and would
soon be flowering. A visit was made to the area on 7 May. Thousands of tulips were observed in full bloom on
9 May 2009 on the north- and west-facing stony serpentine slopes just a hundred metres from the locality first
noted the previous year. The yellow form (Fig. 2B) was far more abundant than the red (Fig. 2A). It was
obvious that the plants were quite different from T. australis that was also found in the same area (Fig. 1A).
On a third visit to Kukësi district on 14 May 2010, the red-flowered form had finished flowering and the
yellow form was already in early fruit, unlike the situation the previous year.

FIGURE 1. Tulipa australis: A: from Surroj area, serpentine substrate. B: from Mt Çajupi, limestone substrate.

The affinities of the new tulip probably lie with the similar T. scardica Bornmüller (1923: 199), which is
restricted to the western Balkan countries of Macedonia and Kosovo (Bornmüller 1923, Hayek 1933, Mayer
& Micevski 1970). A visit was made to the Mrasori river (Mirusha region) in Kosovo on 15 May 2010 in
order to study the populations of T. scardica occurring there. The serpentine habitat of T. scardica in the
Mirusha region is similar to that of T. albanica in the Kukësi district and the distance between the two
localities is less than a hundred kilometres. Tulipa schrenkii Regel (1873: 297) has morphologically similar
flowers, but is found further away from Albania, in the steppes or semi-desert areas of the Ukraine, Caucasus,
Central and Southwest Asia (Botschantzeva 1982, Ghahreman et al. 2007).

18 • Phytotaxa 10 © 2010 Magnolia Press SHUKA ET AL.

Materials and methods

A comparison of the new Albanian tulip with Tulipa scardica and T. schrenkii was carried out based on
literature, herbarium material deposited at Skopje (SKO) and Pristina (herb. Pristina) as well as populations of
T. albanica and T. scardica studied in their natural habitat. All measurements of T. albanica are based on
living material, the mean of 50 plants examined.
Chromosome counts were made on somatic metaphases using the squash technique. Root tips were
obtained from six plants grown from seed collected in the locus classicus. Samples were pretreated with 0.05
% colchicine solution for 5 hours, then fixed in 3/1 (v/v) ethanol/acetic acid for at least 24–48 hours. The root
tips were hydrolysed in 1N HCl for 12 minutes at 60°C, stained with Schiff’s reagent and squashed in a drop
of acetic carmine. Characterisation of karyotype, determination of centromere position and chromosome type
based on several metaphase plates all follow the nomenclature of Levan et al. (1964).
The total DNA amount was assessed by flow cytometry according to Marie and Brown (1993). Triticum
aestivum cv. Chinese spring (2C = 30.9 pg) was used as an internal standard. Leaves of the internal standard
and T. albanica were chopped using a razor blade in a plastic Petri dish with 600 μl of Galbraith nucleus-
isolation buffer (Galbraith et al. 1983) containing 0.1% (w/v) Triton X–100, 10 mM sodium metabisulphite
and 1% polyvinylpyrrolidone 10,000. The suspension was filtered through a 48 μm nylon mesh. The nuclei
were stained with 75 μg/ml propidium iodide, a specific DNA fluorochrome intercalating dye, used with 15
min RNAse treatment (2.5 U/ml) at 4°C. DNA content of 5,000–10,000 stained nuclei was determined for
each sample using a flow cytometer (model CyFlow SL3, Partec, Munster, Germany). The total 2C DNA
value was calculated using the linear relationship between the fluorescent signals from stained nuclei of the
Tulipa species and the internal standard. The mean was calculated from measurements of samples comprising
three seed germinations of T. albanica and one adult individual.

Taxonomy
Tulipa albanica Kit Tan & Shuka, sp. nov.― Figs. 2–4

Tulipa albanica differt ab T. scardica et T. schrenkii foliis inferioribus valde undulatis et canaliculatis, superioribus
apice longe attenuato-rostrato, tepalorum macula basali nigra absente. Ab T. scardica praeterea differt filamentis
aureis (nec eburneis vel nigrescentibus), capsulis minoribus, seminibus majoribus.

Type:—ALBANIA (Northeast Albania: UTM 34T DM1 and 34T DM3). Kukësi district: 6 km from Kolshi
village to Surroj, N- and W-facing open stony serpentine slopes of Ahmet Xhindit, in clearings of Quercus
pubescens woodland and Buxus sempervirens scrub, 625 m, 42°02’N, 20°20’E, 7 May 2009, flowers yellow to
golden-yellow, Shuka 250 (holotype TIR, isotypes C, LD, private herbaria Kit and Shuka).
Paratype:—loc. ibid., flowers scarlet, 7 May 2009, Shuka 251 (TIR, C, private herbaria Kit and Shuka).
Perennial herb. Bulb ovoid to ovoid-globose, 1.5–2.5 cm in diameter, not stoloniferous; tunics dark
reddish-brown, chartaceous, prolonged into short neck; inner surface densely covered with straight, 2.5–3.5
mm long unicellular hairs. Stem erect, 13–40 cm (including subterranean portion), glabrous, glaucous to
greyish-green. Leaves 3–5, alternate, glabrous, glaucous to greyish-green, subacute to mucronulate.
Lowermost leaves (at ground level) linear-lanceolate to broadly-lanceolate, patent to erecto-patent, 10–25 ×
1–3.5 cm, canaliculate, strongly undulate. Upper cauline leaves much smaller, weakly canaliculate, slightly
undulate, ciliate at base, narrowing and long-tapering at apex. Flowers solitary (very rarely 2), large, erect,
campanulate, existing in two colour forms, yellow to golden-yellow or carmine-scarlet turning deep reddish-
maroon. Perianth segments slightly unequal; outer segments elliptical to obovate, 4–7.3 × 2.3–3.6 cm, obtuse-
or rounded-apiculate; inner segments elliptic-oblanceolate to obovate-spathulate, 3.5–7 × 2–4.2 cm, obtuse to
subacute. Perianth in red-flowered forms with conspicuous yellow zone or blotch extending one-sixth to one-
third the length of a segment, visible outside. Filaments golden-yellow, 7–14(–17) × 1.5–2.5 mm, glabrous,

A NEW TULIP FROM NE ALBANIA Phytotaxa 10 © 2010 Magnolia Press • 19

dilated at base. Anthers burgundy-red to blackish-maroon, oblong, 7–13 × 2–3.5 mm, c. equalling or shorter
than filaments; pollen blackish-maroon. Ovary subsessile, light to yellowish-green, 12–23 mm long; stigmas
3, recurved-decurrent, rose-pink, yellow or pale green. Capsule pale brown, ellipsoid, 3–3.5 × 1.5–2 cm,
apiculate, shortly stipitate, with distinct transverse veins throughout. Seeds flat, triangular-deltoid, 5.5–6.5 ×
4.5–6 mm, greenish-brown, turning dark reddish-brown at maturity. Flowering late April to mid-May;
capsules dehiscing by early July.

FIGURE 2. Tulipa albanica: A: habit. B: flower of yellow form. C: stamens, ovary, stigma of yellow and red-flowered
forms. D: capsule and seeds (enlarged). E: bulb. Bar scale 1 cm.

Comparison with Tulipa scardica and T. schrenkii

The serpentine habitat of Tulipa albanica is similar to that of T. scardica in Radusha (locus classicus in
Macedonia) and T. scardica in Mirusha region in Kosovo. According to Mayer & Micevski (1970), plants of
T. scardica show great variation in several morphological characters such as leaf form, flower colour, length
of filaments and anthers in different parts of its distribution range. Our field and herbarium studies confirm
this variation in and between populations. Thus specimens of T. scardica from Taorska Klisura (collected by
Micevski on 9 May 1964, SKO) and specimens from Mirusha region (collected by Krasniqi on 11 May 2002,
herb. Pristina) have flat, slightly undulate leaves and perianth segments with a faint black basal blotch edged

20 • Phytotaxa 10 © 2010 Magnolia Press SHUKA ET AL.

pale or bright yellow. The filament colour varies from blackish in Radusha (locus classicus) to white or pale
purplish in tulips from Mirusha. During our fieldwork to Mirusha region on 15 May 2010, we have observed
plants without any marking on their perianth segments and plants with white basal markings on their perianth.
This supports the establishment of four colour forms within the T. scardica complex by Mayer & Micevski
(1970). We consider the recently described T. serbica Tatić & Krivošej (1997: 733) as falling within the range
of variation of T. scardica. Specimens from Crveni Bregovi-Orlov rid (collected by Micevski on 20 May
1965, SKO) have undulate leaves and flowers with a distinct black basal blotch within and superficially
resemble red-flowered forms of T. albanica, but the latter can easily be distinguished by the complete absence
of a black basal blotch on its perianth segments. It is noteworthy that golden-yellow flowers, which are
predominant in T. albanica (comprising more than 80% of the population), have never been observed in T.
scardica.
As in Tulipa scardica, variation in the colour of the perianth and stamens has also been observed in T.
schrenkii native to the steppes and semi-desert areas of Turkey, Iran, the Caucasus, Daghestan, Kazakhstan,
Azarbaijan, western Siberia and the Ukraine (Botschantzeva 1982). In T. albanica we have only observed
yellow and red flower forms and the stamen colour is always constant regardless of perianth colour, i.e., the
filaments are always golden-yellow and the anthers dark blackish-maroon (Fig. 2C). Tulipa albanica differs
from T. scardica and T. schrenkii by its canaliculate, strongly undulate lower leaves, the upper leaves with a
long-tapering apex (Fig. 3) and the absence of a black basal blotch on the perianth segments. From T. scardica
it additionally differs by its golden-yellow (not whitish or blackish) filaments, smaller capsules and larger
seeds. Selected morphological characters of Tulipa albanica, T. scardica and T. schrenkii are summarized in
Table 1.

TABLE 1. Morphological differences between Tulipa albanica, T. scardica and T. schrenkii.

Characters T. albanica T. scardica T. schrenkii

Leaf number 3–5 3–5(–7) 3(–4)

Leaf form canaliculate, strongly flat, weakly undulate flat, not or weakly
undulate undulate
Lowermost leaves 10–25 × 1–3.5 cm 10–30(–40) × 1.5–4 cm 8–12 × 1.5–4 cm

Upper leaves 3–11 × 0.5–1 cm 4–12 × 0.5–1 cm 4–6 × 0.8–1.5 cm

Colour of perianth segments yellow or scarlet to dark variable except yellow variable
reddish-maroon

Base of perianth segments yellow, without black yellow or cream with or yellow, usually with
blotch without black blotch black blotch

Length of outer perianth 4–7.3 cm 2.4–5.5 cm 2.3–7 cm

segments

Inner perianth segments 3.5–7 × 2–4.2 cm 3–5 × 1.5–2.5 cm 2.3–6 × 1–2.1 cm

Filament colour and length golden-yellow, white or blackish, yellow or blackish,
7–14(–17) mm 6–9(–11) mm 6–12.5 mm
Anther colour and length blackish-maroon, yellowish or dull violet- yellow or dark purple,
7–13 mm purple, 6–13 mm 5–13 mm
Ovary length 12–23 mm 12–22 mm 10–22 mm

Capsule size 30–35 × 15–20 mm c. 42 × 32 mm 20–40 × 22–30 mm

Seed size 5.5–6.5 × 4.5–6 mm c. 5 × 4 mm –

A NEW TULIP FROM NE ALBANIA Phytotaxa 10 © 2010 Magnolia Press • 21

FIGURE 3. Leaves: A: Tulipa albanica, canaliculate and strongly undulate. B: T. scardica, flat, weakly undulate. C: T.
schrenkii, flat, not or weakly undulate.

Chromosome number and karyotype features

The chromosome number of 2n = 2x = 24 is the first count for Tulipa albanica. This is the characteristic
number for the genus Tulipa, in which most of the cultivated species have been cytogenetically investigated
(Blakey & Vosa 1981, 1982, Mizuochi et al. 2007). The chromosomes are large (5 to 8 μm). Two pairs are
metacentric (pairs V and VIII), three submetacentric (IV, X and XII) and the remaining seven pairs are
subtelocentric (I, II, III, VI, VII, IX and XI). The chromosome pairs X and XII seem to have very small
satellites on the long arms (Fig. 4).

Genome size

In Tulipa albanica the nuclear DNA values are 2C = 54.15 (± 0.23) pg, which represent a smaller genome size
as compared to 2C = 61.5–61.8 pg in T. schrenkii and 2C = 69 pg in T. scardica (Zonneveld 2009). In the
panel of diploid Tulipa species measured by Zonneveld (2009) the smallest genome size was 30.9 pg (T.
clusiana f. cashmeriana) and the largest 70.5 pg (T. gesneriana). Tulipa albanica with 2C DNA values of
54.15 (± 0.23) pg represents the intermediate genome size in this panel. The nuclear DNA amount supports
the dissimilarities of the three taxa and indicates that T. albanica is quite distinct from T. scardica and T.
schrenkii. This is the first genome size estimation for a wild tulip originating from Albania.

22 • Phytotaxa 10 © 2010 Magnolia Press SHUKA ET AL.

FIGURE 4. Tulipa albanica: metaphase plate (2n = 24) and corresponding karyogram.

Habitat and some interesting associated species

The locality of the new tulip is to the left of the Black Drin and the river valley has a Mediterranean-type
climate or at least, is under this influence. The predominant vegetation consists of Quercus pubescens Willd.,
Q. cerris L., Carpinus orientalis Mill. and Fraxinus ornus L. woodland and Buxus sempervirens L. scrub. In
the small area of two hectares where T. albanica occurs, several palaeo-endemics restricted to serpentine areas
in the Balkans are also found. They have a narrow distribution range in Albania and include Forsythia
europaea Degen & Baldacci, Centaurea candelabrum Hayek & Košanin, Aristolochia merxmuelleri Greuter
& E.Mayer, Halacsya sendtneri (Boiss.) Dörfler and Paramoltkia doerfleri (Wettst.) Greuter & Burdet. It is
presumed that T. albanica has a limited range in Albania, but it may possibly also occur in Kosovo. Taxa in
the same locality but with a wider distribution range in Albania are Polygala doerfleri Hayek, Sanguisorba
albanica Andras. & Javorka, Veronica andrasovszkyi Javorka and Aster albanicus (Degen) Degen subsp.
albanicus. The Tertiary relict species Euphorbia glabriflora Vis., Genista hassertiana (Bald.) Bald. ex
Buchegger and Dioscorea balcanica Košanin are also present.

Other specimens examined

Tulipa albanica. ALBANIA: Kukësi district, hill of Polla e Zezë, near Surroj village, 22 May 2008, Shuka &
Hallaçi 235 & 236 (private herbarium Shuka); 6 km from Kolshi to Surroj village, N- and W-facing hill

A NEW TULIP FROM NE ALBANIA Phytotaxa 10 © 2010 Magnolia Press • 23

slopes of Ahmet Xhindit, in Quercus woodland or Buxus scrub, 650 m, 7–9 May 2009, flowers yellow to
golden-yellow, Shuka & al. 252–257 (private herbarium Shuka); loc. ibid., flowers scarlet, Shuka & al. 258–
262 (private herbarium Shuka); loc. ibid., 14 May 2010, flowers yellow to golden-yellow, Shuka & al. 1207–
1210 (private herbarium Shuka); loc. ibid., flowers scarlet, Shuka & al. 1211–1213 (private herbarium Shuka).
Tulipa scardica. MACEDONIA: Taorska Klisura, vo pravec na T. Veles, 9 May 1964, leg./det. Micevski
(type of f. setulosa Mayer & Micevski, 3 specimens in SKO); Zeden supra Radusha, 21 May 1965, leg./det.
Mayer & Micevski (type material of f. umbrosa Mayer & Micevski and f. integra Mayer & Micevski, 4
specimens in SKO); Crveni Bregovi-Orlov rid, 20 May 1965, leg./det. Micevski (2 specimens in SKO);
Klisura na r. Pcinja (Sveti Jovan), 26 April 1967, leg./det. Micevski (2 specimens in SKO). KOSOVO: loc.
ign., 12 May 1991, coll. ign. (herb. Pristina, sub T. boeotica); above Mrasori tributary, Mirusha region, 11
May 2002, leg./det. E. Krasniqi 57–60 (herb. Pristina); loc. ibid., 14 May 2010, leg./det. Shuka 1214–1218
(private herbarium Shuka).
Tulipa schrenkii. KAZAKHSTAN: by Lake Tengiz, 275 m, 1 May 2008, Aanensen (photo); near
Korgaldzin, 100–150 km southwest of Astana, steppe, 275 m, 1 May 2008, Aanensen s.n. (Fig. 2C);
Korgaldzin, 325 m, 1 May 2008, Aanensen (photo).
All specimens collected in Albania and Kosovo are from serpentine substrate. The specimens collected in
Macedonia by Micevski and deposited in SKO are stated to be from serpentine and limestone.

Conclusions

Tulipa albanica represents a new species from the serpentine substrate of Kukësi district (northeastern
Albania). We think that the species may have a wider distribution in the Balkans, possibly occurring in similar
habitats in Kosovo.
The affinities of Tulipa albanica lie with T. scardica, a species restricted to limestone and serpentine
substrates of neighbouring Macedonia and Kosovo. Tulipa albanica is morphologically similar to T. schrenkii,
which is found in the more distant steppes of the Ukraine, the Caucasus, and Central and Southwest Asia.
Tulipa albanica differs significantly from T. scardica and T. schrenkii, not only in morphological features
such as leaf form, colour of perianth segments, colour of filaments and anthers, but also in its total nuclear
DNA amount (2C = 54.15 pg), which is less than in the other two tulip species.

Acknowledgements

We are grateful to Prof. V. Stevanović , Zoran Krivošej and Dr. D. Prodanović (all from Belgrade), H.O.
Aanensen (Hisøy, Norway), Prof. V. Matevski (Skopje) and Dr. E. Krasniqi (Pristina) for their helpful
comments, loan of photographs and/or specimens from their herbaria. Special thanks are offered to Prof.
Manfred A. Fischer (Vienna) for kindly translating the diagnosis to Latin. We are much indebted to Mr. B.
Hallaçi and L. Vata for their assistance during fieldwork in northeastern Albania.

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