Article ISSN 1179-3163 (Online Edition)

Tulipa kosovarica (Liliaceae), a new species of tulip from Kosovo

LULËZIM SHUKA1, KIT TAN2 & ELEZ KRASNIQI3

1Department of Biology, Faculty of Natural Sciences, Tirana University, Bld. ZOG I, Albania 2Institute of Biology, University of Copenhagen, Øster Farimagsgade 2D, DK-1353 Copenhagen K, Denmark. E-mail: [email protected] 3Department of Biology, Faculty of Mathematics and Natural Sciences, Pristina University, Mother Theresa str., Kosovo

Abstract

Tulipa kosovarica (Liliaceae) is illustrated and described as a new species from a serpentine area in Rahoveci province, southwestern Kosovo. It is related to T. serbica from northern Kosovo and had previously been confused with T. scardica from (F.Y.R.) Macedonia and southern Kosovo, all found on serpentine soils. Comparison is made with the recently described T. albanica from northeastern Albania. Ecological data for T. kosovarica is presented in detail and the distribution of all four species in the Balkan Peninsula is mapped.

Key words: Balkan Peninsula, serpentine substrate, taxonomy, Tulipa albanica, T. scardica, T. serbica

Introduction

In spite of its small territorial area, Kosovo has an extremely rich flora, due to its varied climate, topography and geology. The number of vascular plant species is estimated as ca. 1800 (Ministry of Environment and Spatial Planning [MESP] 2010) and comprises diverse floristic elements, in particular Balkan endemics and paleo-endemics. According to MESP, biodiversity ‘hot-spots’ are the Sharri mountains in the southwest, the Prokletije mountains (Bjeshkët e Nemuna) in the northwest, and the Koritniku and Pashtriku mountains, both situated in the south of Kosovo. The genus Tulipa in Kosovo is represented by three native species: the widespread T. australis Link (in Schrader 1799: 317), which some botanists treat as a subspecies of the tetraploid T. sylvestris Linnaeus (1753: 305), T. serbica Tatić & Krivošej (1997: 733) a species first described from Mt. Rogozna and the hill Beli Laz, two localities on the opposite sides of the Ibar river, and T. scardica Bornmüller (1923: 199), which is distributed in the Vardar valley in Macedonia and Krivenik, southern Kosovo, with an outlying locality to the west. In this paper a fourth species known from two localities in the serpentine area of Mirusha in Gjakova district, Rahoveci province in southwestern Kosovo is described. On 14 May 2010, two of the authors (L.S. & E.K.) together with Prof. F. Rexhepi and Prof. F. Millaku from Pristina University, went to the Mirusha area at the foot of Mt Kozniku in Rahoveci province, Kosovo to investigate plants that had been earlier reported by E.K. as T. scardica (Rexhepi & Krasniqi 2004). They also planned to compare the material with the recently described T. albanica Kit Tan & Shuka from the Kukësi district in northeastern Albania, occurring less than a hundred kilometres from this site (Shuka et al. 2010: 19). In mid-May, most of the Mirusha plants had finished flowering and were in early fruit. Some important floral characters were observed but we wanted to be convinced as to their consistency. Therefore L.S., together with Z. Hashani (PhD student at Department of Biology, Tirana University), went to the area again in early May and mid-July 2011 to confirm initial observations. Morphometric measurements were obtained from numerous plants in full bloom and later, in fruit. We are now convinced that the Mirusha plants differ from T. scardica to which it had previously been assigned (Rexhepi & Krasniqi 2004, Shuka et al. 2010) and furthermore, it appears to be more similar to T. serbica.

Accepted by Maarten Christenhusz, 25 Jul. 2012; published online in PDF: 8 Aug. 2012 1 Description of new species

Tulipa kosovarica Kit Tan, Shuka & Krasniqi, sp. nov.― Figs. 1–4

FIGURE 1. Tulipa kosovarica: A, whole plant; B, bulb; C, capsule; D, seeds. Scalebar 1 cm.

2 • Phytotaxa 62 © 2012 Magnolia Press SHUKA ET AL. Distinguished from Tulipa albanica and T. scardica by its white perianth base and white filaments, and from T. serbica by its cerise-magenta, obovate inner perianth segments, which are not pale pink nor rhomboidal in shape, as well as by its clearly demarcated white perianth base. Type:—KOSOVO (Southwestern Kosovo: UTM 34T DN4). Rahoveci province: Mirusha region at the foot of Mt Kozniku, N-facing serpentine slopes on the left side of Mrasori tributary, branch of White Drini river, between Mrasori and Llapçevë villages, ca. 1.5 km above the railroad, in open thermophilous woodland dominated by Quercus pubescens and Juniperus oxycedrus, 420–480 m, 42°30’N, 20°34’E, 20 May 2011, flowers red to scarlet, white at the base, Shuka 4578 (holotype TIR, isotypes C, LD, herb. Pristina, private herbarium Shuka).

FIGURE 2. Flowers: A & B, Tulipa kosovarica; C, T. albanica; D, T. scardica. Scalebar 1 cm.

Perennial herb. Bulb 3–4 × 1.5–2.0 cm, ovoid, not stoloniferous; tunics blackish-brown, chartaceous; inner surface densely covered with straight, 2.5–4.0 mm long silky adpressed unicellular basifixed hairs. Stem erect, 15–30 cm above ground (excluding subterranean portion), glabrous, glaucous to greyish-green. Leaves 3–5, lower ones alternate, upper two usually opposite, glaucous to greyish-green, erect-patent, not exceeding

TULIPA KOSOVARICA, A NEW SPECIES OF TULIP FROM KOSOVO Phytotaxa 62 © 2012 Magnolia Press • 3 the flower, entire; lowermost leaves (at ground level) 10–26 × 1.5–3.0 cm, broadly canaliculate, weakly undulate, linear-lanceolate, apiculate to subacute, cucullate when young, ciliolate in lower 2/3; upper cauline leaves 5–11 × 0.5–0.8 cm, smaller and narrower, acute to acuminate. Flowers solitary, erect, campanulate, glabrous. Perianth segments slightly unequal, cerise to dark magenta-red with white apiculate papyraceous tips and white to whitish base occupying the lower quarter, the white base less visible on external surface of perianth segments and not conspicuous at all in dried material. White base sometimes suffused by the reddish- purple colour of the perianth segments, resulting in obtrullate blackish-violet basal patches or dark shadows. Outer segments 3.5–5.0 (–6.0) × 1.5–2.3 cm elliptic, attenuating to a narrow claw at base; inner segments 3.0–4.7 × 1–2 cm, obovate to oblanceolate, obtuse to subacute. Filaments 10–13 mm, white, glabrous, dilated at base; anthers 8–12 mm, blackish-maroon, oblong to elliptical, shorter than filaments; pollen purplish- maroon to dull purple at anthesis, very rarely yellow, olivaceous with age. Ovary subsessile, 15–20 mm long (longer than filaments at anthesis), yellowish-green; stigmas 3, recurved-decurrent, pale green, papillose. Capsule 3.0–3.5 × 1.5–2.2 cm, ellipsoid, apiculate, shortly stipitate, with distinct transverse veins throughout, pale brown. Seeds 7–8 × 6 mm (including the 0.8–1.2 mm broad wing), flat, triangular-deltoid; brown, turning dark reddish-brown at maturity. Flowering late April to mid-May; capsules dehiscing end of June to mid-July.

FIGURE 3. A & B, flower and rhomboidal inner perianth segment without white base of T. serbica; C, obovate inner perianth segment with white base of T. kosovarica. Scalebar 1 cm.

Tulipa kosovarica (Figs. 1 & 2A–B) differs from T. scardica by its white or whitish perianth base that is sometimes masked by obtrullate patches of maroon and violet. However, the perianth base is never yellow with blackish-purple blotches as is typical for T. scardica (Fig. 2D). An important difference lies in the stamen colour: the filaments of T. kosovarica are always white and the anthers and pollen maroon to dark purple, very rarely yellow. In T. scardica the filaments are never white but dark-coloured, the anthers are blackish-maroon and the pollen greenish-yellow at dehiscence (Fig. 4D), very rarely blackish-maroon. From T. serbica, the most obvious difference is in the shape and colour of the inner perianth segments and colour of the pollen grains. The inner perianth segments in T. kosovarica are obovate (Fig. 3C) and cerise-magenta (Fig. 2A–B), but in T. serbica they are more rhomboidal in shape (Fig. 3B) and pale pink with a darker pink broad central band so that the segment appears pink with broad white margins (Fig. 3A). The base of the perianth in T. serbica is usually pale pink within and not demarcated in colour from the rest of the perianth; it is rarely

4 • Phytotaxa 62 © 2012 Magnolia Press SHUKA ET AL. whitish to pale green and never with black blotches. The pollen is yellow (Fig. 4B), very rarely purplish- maroon. White-flowered individuals exist in the wild as noted by Prodanović and Krivošej (Stevanović, pers. comm. 2009). Tulipa albanica differs from the other three named species by its combination of yellow perianth bases without black blotches, yellow filaments (Fig. 2C) and violet-purple pollen (Fig. 4C). The morphological differences in the four species are summarized in Table 1.

TABLE 1. Comparison of morphological characters in Tulipa albanica, T. kosovarica, T. scardica and T. serbica. Characters Tulipa albanica Tulipa kosovarica Tulipa scardica Tulipa serbica Leaf number 3–5 (2–)3–4 (–5) 3–5 3–5 Lowermost leaf canaliculate, strongly broadly canaliculate, broadly canaliculate, canaliculate, weakly undulate weakly undulate weakly undulate undulate Lowermost leaves 10–25 × 1.0–3.5 cm 10–22 × 1.5–3.0 cm 10–20 × 1.5–2.5 cm 10–30 × 1.5–3.0 cm Leaf margin ciliolate in basal 1/3 ciliolate in basal 2/3 ciliolate in basal 1/3 or ciliolate throughout 2/3 Upper leaves 3–11 × 0.5–1.0 cm 5–12 × 0.5–0.8 cm 4–12 × 0.5–1.2 cm 3–6 × 0.6–0.8 cm Colour of perianth yellow or scarlet to cerise-magenta scarlet to burgundy red white or pale pink segments dark reddish-maroon edged white at margin Base of perianth yellow, without black white or whitish without yellow or cream with a pale pink or whitish to segments blotch black blotch, sometimes blackish-purple blotch pale green, without masked by shades of black blotch maroon and purple Length of outer 4.0–7.3 × 2.3–3.6 cm 3.5–5.0 (–6.0) × 1.5–2.3 cm 3.5–4.5 × 1.3–2.1 cm 2–5 × 1.2–1.8 cm perianth segments Inner perianth 3.5–7.0 × 2.0–4.2 cm 3.0–4.7 × 1–2 cm 3–5 × 1.5–2.5 cm 2.5–4.5 × 1.2–2.0 cm segments Filament colour golden-yellow, white, 10–13 mm blackish-purple in upper white or whitish- and length 7–14 (–17) mm half, yellow in basal coloured, 6–7 (–9) mm half, 6–9 (–11) mm Anther colour and burgundy-red to blackish-maroon, 8–12 mm blackish-maroon, maroon to dull purple, length blackish-maroon, 7–9 mm 6–9 mm 7–13 mm Pollen colour at dark violet-purple, purplish-maroon, very greenish-yellow, very yellow, very rarely dehiscence very rarely yellow rarely yellow rarely blackish-maroon purplish-maroon Ovary length at 12–23 mm 15–22 mm 20–22 mm 9–15 mm anthesis Stigma colour rose-pink, yellow or cream to pale green pale yellow cream to pale green pale green Capsule size 30–35 × 15–20 mm 30–35 × 15–22 mm 34–44 mm long 20–30 × 15–20 mm Seed size 5.5–6.5 × 4.5–6.0 mm 7–8 × ca. 6 mm ca. 5 × 4 mm immature

Among the Balkan tulips with non-woolly inner bulb tunics and glabrous filaments (Tulipa sect. Tulipa), there is the frequent combination of red perianth segment with black basal blotches edged yellow, together with greenish-yellow (as in T. scardica) or blackish-maroon pollen grains (as in T. rhodopea (Velen.) Velen.); this syndrome is exhibited also in T. undulatifolia Boiss. that belongs, however, to a different section, T. sect. Tulipanum Reboul. This set of characters is absent in T. kosovarica, which has a combination of red perianths with white bases, absence of black basal blotches, presence of white filaments and predominantly purplish- maroon pollen grains (Fig. 4A), thus differing from the other species in T. sect. Tulipa. Also in this respect, T. serbica with its combination of white or pale-pink perianth, absence of black basal blotches, presence of white or whitish-coloured filaments and predominantly yellow pollen (Fig. 4B) deserves taxonomic recognition from T. scardica to which it had previously been assigned (Shuka et al. 2010). Observations made on several

TULIPA KOSOVARICA, A NEW SPECIES OF TULIP FROM KOSOVO Phytotaxa 62 © 2012 Magnolia Press • 5 taxa in Greece indicate that the combination of black basal blotches on a yellow background together with red perianths is a reliable and easily observed distinguishable character. Pollen colour alone is not a distinguishing feature between species, but is useful when studied in combination with several other characters. Tulipa scardica with a nuclear DNA content of 2C = 69 pg (Zonneveld 2009) is treated as a synonym of T. gesneriana Linnaeus (1753: 306) in the World Checklist for Tulipa (Govaerts 2012); the latter species has a 2C value averaging 68.7 pg, which is also close to that of 2C = 68.6 pg for T. rhodopea (Zonneveld 2009). Tulipa undulatifolia Boiss., which has similar perianth markings is however, in a different section, and this taxonomic distinction is represented by its much lower 2C value of 51.2 pg (Zonneveld 2009).

FIGURE 4. Pollen grains: A, T. kosovarica (purplish-maroon); B, T. serbica (yellow); C, T. albanica (dark violet-purple); D, T. scardica (greenish-yellow). Scalebar 0.5 mm.

Tulipa kosovarica was discovered growing on serpentine substrate in only two localities in southwestern Kosovo (Fig. 5), in the watershed of the White Drin, one of the two main branches of the Drini river that flows into the Adriatic Sea. The climate is of a Mediterranean type, with warm, dry summers and cool, wet winters, the average annual temperature ranging from 9º to 13ºC (Avdullahi et al. 2007). The average precipitation in the Mirusha watershed varies from 600 to 800 mm annually. Rainfall is irregular, with 70 to 80% falling between October and the end of April. Therefore plants flowering in spring continue their life cycle in semi- drought. The soils are generally poor in nutrients (Shuka et al. 2009) and support during early summer a low herbaceous cover with a high evaporation coefficient. Our observations carried out during two years indicate that the degree of leaf undulation in plants of T. kosovarica and T. albanica depends considerably on soil moisture content and sun exposure. Plants growing in shaded or densely vegetated places have less undulate leaves than plants growing on open, sunny, bare stony ground.

FIGURE 5. Distribution of Tulipa albanica, T. kosovarica, T. scardica and T. serbica in the Balkan Peninsula.

TULIPA KOSOVARICA, A NEW SPECIES OF TULIP FROM KOSOVO Phytotaxa 62 © 2012 Magnolia Press • 7 The habitat of T. kosovarica in the upper White Drin river is almost identical to that of T. albanica in the lower Black Drin river valley in Albania, and both areas are rich in relict and endemic taxa. The floristic composition of the habitats, where T. kosovarica and T. albanica occur, includes several palaeo-endemics such as Alyssum markgrafii O.E.Schulz, Aristolochia merxmuelleri Greuter & E.Mayer, Aster albanicus Degen subsp. albanicus, Euphorbia glabriflora Vis., Forsythia europaea Degen & Bald., Genista hassertiana (Bald.) Buchegger, Halacsya sendtneri (Boiss.) Dörf., Haplophyllum boisserianum Vis. & Pančić, Minuartia baldaccii (Halácsy) Mattf. subsp. baldaccii, Paramoltkia doerfleri (Wettst.) Greuter & Burdet and Sedum serpentini Janch. These taxa are obligate serpentinophytes, restricted to serpentine areas in the Balkans. Non palaeo-endemics (not solely restricted to serpentine) include Acer tataricum L., Crocus dalmaticus Vis., Dictamnus albus L., Fraxinus ornus L., Iris reichenbachii Heuff., Orchis ustulata L., Linum hirsutum L., Potentilla heptaphylla L. subsp. australis (Nyman) Gams var. malyana (Borbás ex Malý) G.Beck, Rorippa pyrenaica (All.) Rchb. and Tulipa australis Link. Rare taxa associated with the T. kosovarica habitat, but not found at the T. albanica sites are Allium meteoricum Halácsy, Centaurea kosaninii Hayek, Potentilla visianii Pančić, Saponaria sicula Raf. subsp. intermedia (Simmler) Chater, Serratula radiata (Waldst. & Kit.) M.Bieb. subsp. cetinjensis (Rohlena) Hayek and Stipa mayeri Martinovský. Taxa associated with the T. albanica habitat and not found at the T. kosovarica sites are Centaurea candelabrum Hayek & Košanin, Cerastium grandiflorum Waldst. & Kit. subsp. serpentini F.K.Mey., Chamaecytisus purpureus (Scop.) Link, Crocus biflorus Mill. subsp. weldenii (Hoppe & Fürnr.) K.Richt., Dioscorea balcanica Košanin, Echium russicum J.F.Gmel., Fritillaria montana Hoppe ex W.D.J.Koch, Gypsophila spergulifolia Griseb., Lilium albanicum Griseb., Polygala doerfleri Hayek, Sanguisorba albanica András. & Jáv. and Veronica andrasovszkyi Jáv. Tulipa serbica was first discovered by Pavlović on Mt Rogozna but then misidentified as Bornmüller’s T. scardica (Pavlović 1962). It was later collected by Tatić & Krivošej (1997) from Beli Laz hill near the village of Srbovac, a locality near Mt Rogozna, on the opposite side of the Ibar river that flows into the Danube; here it inhabits open stony pastures in Quercus frainetto Ten. woodland (Tatić & Krivošej 1999). Two additional localities with ca. 2000–3000 individuals were discovered more recently by Danijela Prodanović above the village of Lozište in the Ibar Gorge (Prodanović et al. 2008). Both T. kosovarica and T. serbica are probably local endemics with restricted distributions in southwestern and northern Kosovo respectively. Tulipa scardica has a wider distribution, occurring on both sides of the Vardar watershed with an outlying locality to the west at Bistra (Fig. 5). The Vardar river flows southwards into the Aegean Sea. The southern and lower areas of the watershed are dominated by Quercus frainetto, and the middle and upper regions covered with Carpinus orientalis Mill. The most abundant species in these localities are Achillea fraasii Schultz Bip., Aristolochia macedonica Bornm., Carex nitida Host., Fraxinus ornus L., Helianthemum canum (L.) Baumg., H. nummularium (L.) Mill., Iris reichenbachii Heuff., Juniperus excelsa M.Bieb., Minuartia glomerata (M.Bieb.) Degen, Orchis tridentata Scop., Polygala vulgaris L., Syringa vulgaris L. and Vicia onobrychoides L. (Mayer & Micevski 1970). These are non-facultative serpentinophytes, occurring on both serpentine and limestone soils in the open clearings of trees or scrub.

Additional specimens examined

Tulipa kosovarica. KOSOVO: Mirusha region, N-facing serpentine slopes on the left side of Mrasori tributary, in open vegetation, 420–480 m, 42°30’N, 20°34’E, 11 May 2002, flowers scarlet, Krasniqi 57 (herbarium, Faculty of Mathematics and Natural Sciences, Pristina University); loc. ibid., 15 May 2003, Krasniqi (photos); loc. ibid., 14 May 2010, Shuka 1214 (TIR, ATH, private herbaria Kit and Shuka); loc. ibid., 14 May 2010, Shuka 1215 (private herbarium Shuka); loc. ibid., 20 May 2011, Shuka 4579 & 4580 (private herbarium Shuka); cultivated material at Copenhagen originating from bulbs collected in locus classicus, photographed and pressed 1 May 2012, Kit Tan & G. Vold (C); near village of Guriq (Malishevë), NE-facing serpentine slopes, 400–500 m, 7 May 2012, Krasniqi, Millaku & Rexhepi 1125–1129 (private herbarium Krasniqi).

8 • Phytotaxa 62 © 2012 Magnolia Press SHUKA ET AL. Tulipa scardica. MACEDONIA: Taorska Klisura, vo pravec na T. Veles, 9 May 1964, Micevski (type of f. setulosa Mayer & Micevski, 3 specimens in SKO); Zeden supra Radusha, 21 May 1965, 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, Micevski (2 specimens in SKO); Klisura na r. Pcinja (Sveti Jovan), 26 April 1967, Micevski (2 specimens in SKO); [ditionis Scardicae in collibus aridis saxosis (substratu serpentine)] prope Raduše [ad basin meridionalem montium catenae Šar-dagh], ca. 400 m, 28 April 1918, Bornmüller Plantae Macedoniae no. 5002 (isotype in B, digital specimen image); in declivitat. prope Raduše (inter Usküb et Totovo), 400–500 m, 28 April 1918, Bornmüller Plantae Macedoniae no. 5002 (lectotype in B, digital specimen image); Negotino to Lakavica, Konecka plain, cultivated material at Göteborg Botanical Garden, accession no. SZ88- 060; Bistra, cultivated material at Göteborg Botanical Garden, accession no. 1991-0540.

Tulipa serbica. KOSOVO: hill of Beli Laz, serpentine slopes, 800–900 m, May 1992, coll. ign. (5 sheets in herbarium, Faculty of Mathematics and Natural Sciences, Pristina University); Mt Rogozna, near Donja Kamenica, serpentine slopes, 900 m, Prodanović (photos).

Acknowledgements

We thank Professors F. Rexhepi and F. Millaku (Pristina University, Kosovo) and Z. Hashani (Tirana University, Albania) for accompanying us during fieldwork and for their kind assistance. We are grateful to Prof. V. Stevanović (Belgrade, Serbia) for making available photographs of Tulipa serbica taken at the locus classicus by Dr D. Prodanović (Zubin Potok University, Kosovo) and to Dr D. Prodanović for sending us additional photographs from new localities. Prof. A. Strid (Ørbæk, Denmark) kindly provided Fig. 1D (T. scardica), which was photographed from cultivated material at the Göteborg Botanical Garden and Dr S. Siljak-Yakovlev (Orsay, France) is thanked for her comments on an earlier draft.

References

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