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).
Turk J Bot
30 (2006) 455-458
© TÜB‹TAK
455
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
Research Note
* E-mail: zarre@khayam.ut.ac.ir
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 0.5-
5.5 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
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)1.1-
2(-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 3.5-
5(-7) mm wide. Seeds light brown, narrowly oblong, 2.3-
3.5(-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
On the Circumscription of Isatis tinctoria L. (Brassicaceae) in Iran
456
25°N
30°N
35°N
40°N
45°E 50°E 55°E 60°E
IRAN
Figure 1. Map of the distribution Isatis tinctoria in Iran.
(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, 1300-
1500 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
H. MOAZZENI, S. ZARRE
457
Table 1. Diagnostic characters useful in distinguishing Isatis tinctoria from its closest relatives in Iran.
Taxon → Isatis gaubae Isatis pachycarpa I. tinctoria
Character↓
Shape of the fruit oblong-linear obovate-elliptic obovate to elliptic, rarely oblong
Carinate fruit locule absent present absent
Width of the fruit locule 2-3 mm 3 mm 1.5-2 mm
Width of the wings at the middle of the fruit 1-1.5 mm 2.5 mm 2-3 mm
1
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.
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.
On the Circumscription of Isatis tinctoria L. (Brassicaceae) in Iran
458
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