Turk J Bot 30 (2006) 455-458 © TÜB‹TAK Research Note On the Circumscription of Isatis tinctoria L. (Brassicaceae) in Iran Hamid MOAZZENI, Shahin ZARRE* Research Centre of Plant Sciences, School of Biology, University College of Science, Tehran University, PO Box: 14155-6455, Tehran - IRAN Received: 05.12.2005 Accepted: 21.06.2006 Abstract: In the course of preparing a revision of Isatis L. and its allies in Iran, we verified Isatis tinctoria L. among the material studied. Although it has been previously recorded for Iran, we give a more extensive description and a map of its distribution in Iran. A key to distinguishing this species from its allies and a discussion of the affinities in this group are provided. Key Words: Brassicaceae, Isatis tinctoria, Iran Introduction Isatis L., with about 50 species (Appel & Al-Shehbaz, 2003), is an extremely difficult genus of Brassicaceae (Cruciferae), which is distributed primarily in the northern hemisphere and has an Irano-Turanian distribution pattern (Davis, 1965). According to Hedge (1968) and additions made by Sajedi et al. (2004), Iran, with 19 species, is one of the important diversification centres of the genus. Extractions of I. tinctoria L. are widely used as the Old World blue dye, woad, which became the dominant blue dye in Europe, especially in western Europe. In some parts of the world, it is a noxious weed, and nowadays it is rarely cultivated. Isatis tinctoria L. (woad) is distributed in south-west Asia, Uzbekistan, Russia, Tajikistan, Mongolia, Pakistan, Korea, Kazakhstan, Japan (Shu, 2001), and Europe (Tan, 2002), and is naturalised elsewhere (e.g., North America). In the recently published Flora Hellenica, 2 subspecies have been considered for I. tinctoria (Tan, 2002), while in the Flora of Turkey and the East Aegean Islands, the species is composed of 4 subspecies (Davis, 1965). There are 2 references recording Isatis tinctoria from Iran. Parsa (1951) recorded this species from Kermanshah province in western Iran based on a specimen collected near Tagh-e Bostan by Cowan and Darlington. Although this specimen was not seen by us, all other specimens identified by Parsa as I. tinctoria and deposited in the herbarium of the Pharmacological Faculty of Tehran University were actually I. kotschyana Boiss. Another specimen verified and published recently as I. tinctoria by Sajedi et al. (2004) from northern Tehran and deposited in IRAN herbarium, is only a form of I. gaubae. Therefore, the presence of I. tinctoria in Iran seemed to be doubtful; however, while preparing a revision of Isatis for the Flora of Iran, we found several specimens, distributed mainly in eastern Iran, which are indeed I. tinctoria. Materials and Methods This study was primarily based on material deposited in the main herbaria of Iran, i.e. FUMH (Ferdowsi University of Mashhad Herbarium), IRAN, TUH, the local herbarium of Kerman University, and the Natural History Museum of Vienna (W). Several field studies were also conducted in north-east Iran. The specimens were compared with the material deposited in W and identified as I. tinctoria by Davis and Hedge (Edinburgh). It is described below and a distribution map for Iranian specimens is provided (Figure 1). * E-mail: zarre@khayam.ut.ac.ir 455 On the Circumscription of Isatis tinctoria L. (Brassicaceae) in Iran 40°N 35°N IRAN 30°N 25°N 45°E 50°E 55°E 60°E Figure 1. Map of the distribution Isatis tinctoria in Iran. Results and Discussion Taxonomic account Isatis tinctoria L., Sp. Pl. 2: 670. 1753. Type: "Hab. ad littora maris Balthici et oceani Europae", Hb. Linn. 848/1 (microfiche!). Common names: woad, dyers woad. Short-lived perennial (according to Davis, 1965; Hedge, 1968; Tan, 2002), biennial, or rarely, annual herbs, glaucous, glabrous, or pubescent below. Stems (30-)40-100(-150) cm, erect, often paniculately branched above middle. Basal leaves rosulate, petiole 0.55.5 cm; leaf blade oblong or oblanceolate, (2.5-)5-15(20) × (0.5-)1.5-3.5(-5) cm, base attenuate, margin entire, repand, or dentate, apex obtuse; cauline leaves sessile; leaf blade oblong or lanceolate, rarely linearoblong, base sagittate or auriculate, margin entire, apex acute. Fruiting pedicels slender, considerably thickened, and subclavate at apex, 5-10 mm. Sepals oblong, 1.5-2.8 × 1-1.5 mm, glabrous or sparsely pubescent; petals yellow, oblanceolate, 2.5-4 × 0.9-1.5 mm, base 456 attenuate, apex obtuse; filaments 1-2.5 mm; anthers oblong, 0.5-0.7 mm. Fruits black or dark brown, oblongoblanceolate, elliptic-obovate, or rarely oblong, (0.8)1.12(-2.7) cm × 3-7 mm, often broad above middle, glabrous or pubescent, winged all around, base cuneate, margin sometimes slightly constricted, apex subacute, rounded, or rarely subemarginate; loculus with a distinct mid vein and inconspicuous lateral vein; apical wing 3.55(-7) mm wide. Seeds light brown, narrowly oblong, 2.33.5(-4.5) × 0.8-1 mm. 2n = 14, 28. Flowering: April-June. Habitat: Roadsides, field pastures, hillsides, prairies, railroad embankments, waste places, 300-2200 m; south-west Asia, Europe; naturalised in North and South America. Specimens examined (Figure 1): Prov. Kerman: Kerman, 24.6.1976, M. Mousavi & B. Tehrani 15441 (IRAN!); NE Jebale Barez, Jebale Barez, 29.3.2005, M. Mirtajedini 35785 (TUH!); Kerman, Laleh Zar maountain, 26.4.1997, M. Mirtajedini 35827 H. MOAZZENI, S. ZARRE (TUH!). Prov. Khorassan: NE Bojnourd, Darband to Gifan, 1050 m, 1.5.2005, H. Moazzeni 35803 (TUH!); NE Bojnourd, Tang-e Turkaman, 988 m, 3.4.2005, H. Moazzeni & A. Ghorbani 35779 (TUH!). Prov. Gorgan: Golestan national park, pass between Robat-e Gharabil and Solgerd, 5.6.1987, A. Ghahreman & V. Mozaffarian 5884 (TUH!); Birjand, Tannak to Esfazar, 2000 m, M. Faghihnia & H. Zangooie 30268 (FUMH!); Prov. Golestan: Shahpasand to Bojnord, Almeh road, 13001500 m, 7.6.1975, F. Termeh 15649 (IRAN!). Note: Isatis gaubae and I. pachycarpa are most closely related to I. tinctoria. Both of these species are endemic to Iran. From the former, I. tinctoria can be distinguished by its obovate fruits with the wider part at the upper third of the fruit, while the fruits in I. gaubae are more or less linear-oblong. Moreover, in the middle of the fruits, the locules are wider than the wings in I. gaubae, but the fruit wings are distinctly wider than the locules in I. tinctoria. Isatis pachycarpa, known from southern Iran (Jebal-e Barez mountain range), differs from I. tinctoria in that it has distinctly hairy leaves and velutinous hairy fruits. Furthermore, the fruits of I. pachycarpa are distinctly carinate at the locules. Table 1 shows some characteristic features useful in separating the species in this group. Based on the map presented in Figure 1, Isatis tinctoria covers a much wider area in Iran than previously reported. The specimen reported by Sajedi et al. (2004) is distinctly perennial1, with fruit that are wide in the middle and fruit locules are wider than the wings (appropriate with I. gaubae). This specimen collected from central Alborz in northern Tehran in an area adjacent to Dizin is only a form of I. gaubae. According to our concept, I. tinctoria is distributed in eastern Iran and has a wide distribution throughout the rest of Iran. Most probably, the species penetrates Iran from neighbouring countries to the east, i.e. Afghanistan and Pakistan. Isatis tinctoria has also been recorded from Turkey, most eastern up to C9 Hakkari (Davis, 1965), doubtfully from Iraq (Hedge & Lamond, 1980), Caucasus and Central Asia (Vasil'chenko, 1939), Afghanistan (as I. koelzii by Rechinger, 1958; Hedge, 1968), and west of Pakistan (Jafri, 1973). Therefore, its occurrence in Iran is not unexpected. We think that this species is widely distributed in eastern Iran and in adjacent neighbouring countries, i.e. Afghanistan and Pakistan, but based on the map presented in Figure 1 it seems that there are disjunctions in distribution of this species in eastern Iran. The species is recorded from eastern Afghanistan and the Swat/Chitral area in Pakistan, but not from areas adjacent to Iran. However, the borders between Iran, Afghanistan, and Pakistan have been extremely dangerous for long time, making it difficult for botanists to work. More intensive collection in the future may fill the gaps in the distribution of I. tinctoria in this area. Isatis tinctoria is one of the most polymorphic species of the genus and has been cultivated since ancient times as a source of blue dye (woad), which is obtained by fermenting the ground leaves and lower portions of the plant. Yıldırımlı (1988) reported that I. tinctoria had a global dispersion, except for southern Asia, and Central and southern Africa. He also stated that up to 20 species of Isatis could be reviewed under the I. tinctoria complex. Several subspecies and varieties have been recognised in Table 1. Diagnostic characters useful in distinguishing Isatis tinctoria from its closest relatives in Iran. Taxon → Character↓ Isatis gaubae Isatis pachycarpa I. tinctoria Shape of the fruit oblong-linear obovate-elliptic obovate to elliptic, rarely oblong Carinate fruit locule Width of the fruit locule Width of the wings at the middle of the fruit 1 absent present absent 2-3 mm 3 mm 1.5-2 mm 1-1.5 mm 2.5 mm 2-3 mm We did not observe any perennial specimens of Isatis tinctoria, but according to Hedge (1968), Davis (1965), and Tan (2002) this species can also be a short-lived perennial. 457 On the Circumscription of Isatis tinctoria L. (Brassicaceae) in Iran it. Davis (1965) recognised 3 subspecies under this species for Turkey. Hedge (1968) considered I. koelzii as a species separate from I. tinctoria restricted to east Afghanistan and Pakistan, on the basis of its short (ca. 15 mm long) oblong fruits, which are rounded at both ends. However, these characters show considerable variation among specimens of I. tinctoria and cannot be considered diagnostic for separation of taxa at a specific level. For these reasons Jafri (1973) considered it as I. tinctoria subsp. koelzii (Rech.f.) Jafri. Based on some overlapping characters, Tan (2002) also considered 2 subspecies for I. tinctoria in the area covered by Flora Hellenica: subsp. tinctoria and the localised subsp. athoa (Boiss.) Papan. None of the above-named treatments provide an exact definition for the different subspecies. Based on Iranian samples of I. tinctoria, we were not able to recognise subspecies. For example, in one population we observed some individuals matching the type of I. tinctoria subsp. tinctoria and some in accordance with I. tinctoria subsp. koelzii. Even in one specimen, different sizes and shapes were observed. We were also unable to use the concept applied in the recent volume of the Flora of Turkey (Güner et al., 2001) for delimitation of the 4 subspecies of the Iranian material known in that area. Therefore, it seems that the subspecific concept in I. tinctoria should be re-assessed and more precisely defined. A more detailed biosystematic study in the future may clarify the final systematic position of this species along with other taxa related to it. Acknowledgements We are grateful to Dr. Ihsan Al-Shehbaz (Missouri Botanical Garden) for confirming the identifications and his valuable suggestions. We thank Mr. M. Mirtajedini (Kerman University, Iran), who provided 2 specimens from southern Iran, and Mrs. S. Sajedi (IRAN herbarium),who provided the material deposited in IRAN herbarium. References Appel O & Al-Shehbaz IA (2003). Cruciferae. In: Kubitzki K (ed.) Families and Genera of Vascular Plants, Vol. 5: 75-174. Berlin and Heidelberg: Springer-Verlag. Parsa A (1951). Flora de l' Iran, Vol. 1, pp. 853-866. Tehran: Imprimerie Danesh. Rechinger KH (1958). Cruciferae. In: Køeie M & Rechinger KH (eds.) Symbolae Afghanicae IV. Biol Skr Dan Vid Selsk 10: 13-54. Davis PH (1965). Cruciferae. In: Davis PH (ed.) Flora of Turkey and the East Aegean Islands, Vol. 1: 1-567. Edinburgh: Edinburgh University Press. Sajedi S, Sharifnia F & Sonboli A (2004). Verification of Isatis tinctoria in Iran. Rostaniha 5: 51-52. Güner A, Özhatay N, Ekim T & Baser KHC (2001). Flora of Turkey and the East Aegean Islands, Vol. 11 (Supplement 2). Edinburgh, Edinburgh University Press. Shu SL (2001). Isatis. In: Taiyan Z., Lianli L., Guang Y., Al-Shehbaz IA (eds.) Flora of China, Vol. 8: 36-38. Missouri, Missouri Botanical Garden. Hedge IC (1968). Cruciferae. In: Rechinger KH (ed.) Flora Iranica, Vol. 57: 1-372. Graz: Akademische Druck-u. Verlagsanstalt. Hedge IC & Lamond JL (1980). Brassicaceae. In: Townsend CC & Guest E (eds.) Flora of Iraq, Vol. 4 (2): 827-1199. Ministry of Agriculture, Baghdad. Jafri SMH (1973). Brassicaceae. In: Nasir E & Ali SI (eds.), Flora of West Pakistan, No: 55: 1-308. Karachi: University of Karachi. 458 Tan K. (2002). Isatis. In Strid A. & Tan K. (eds.) Flora Hellenica, Vol. 2: 126-127. Ruggell, Gantner Verlag. Vasil'chenko IT (1939). Cruciferae. In: Komarov, V.L. (ed.) Flora of the U.S.S.R., Vol. 8: 13-515. Moskva, Leningrad: Akademii Nauk SSSR.[in Russian] Yıldırımlı S (1988). The revision of the genus Isatis L. (Cruciferae) in Western half and North of Turkey. DOGA TU J Botany 12: 332400.