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Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

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Istanbul Ecz. Fak. Derg. / J. Fac. Pharm. Istanbul
44(1) 2014 s.31-62

Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

Mehmet Bona
Department of Pharmaceutical Botany, Faculty of Pharmacy, Istanbul University,
34116 Istanbul, Turkey

Abstract: This taxonomic revision is based on morphological data that come

from a wide range of herbarium material and living material collected from wild.
The following are provided for each taxon: a description, along with the com-
plete synonymy, geographical distribution, habitat, specimen examined, and pho-
tographs. Lepidium pumilum and L. cartilagineum are recognized at species rank.
Type specimen is designated for L. pumilum. This study also shows that L. pin-
natifidum does not occur in Turkey, although previously it had been considered a
new record for Flora of Turkey. This study recognises 14 Lepidium s.s. taxa from
Turkey. L. virginicum is naturalized invasive alien plant.
Key words: Brassicaceae, Lepidium, revision, Turkey

*Correspondence: [email protected]
32 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

Introduction
The family Brassicaceae comprises 49 tribes, 321 genara and 3660 spe-
cies (1). It also has major scientific and economic importance (2). The first
comprehensive treatment of the family was that of De Candolle (3), who
based his classification on fruit type (longer than wide vs. wider than long)
and seed embryos (position of the radicle in relation to cotyledons in the
seed). Schulz (4) proposed a new tribal classification of the family. Schulz
relied heavily on fruit characters and seed morphology to delimit tribes
and subtribes (5). Brassicaceae has 555 species and 91 genera according
to Flora of Turkey the East Aegean Islands which followed the Schulz’s
(4) classification (6, 7, 8). Recent substantial advances in the molecular
phylogenetic studies on the Brassicaceae (9, 5, 10) have led to numerous
changes in generic delimitations. According to recent classification Turkey
is a centre of diversity with 660 taxa belonging to 91 genera, including
571 species, 65 subspecies and 24 varieties. The taxa of the East Aegean
Islands are excluded from the total species number of Turkey (11).
The genus Lepidium L. is one of the largest genera of the Brassicace-
ae consisting of 250 species (1). It is distributed worldwide, primarily in
temperate and subtropical regions. The genus grows in the mountains in
tropical areas and it is poorly represented in Arctic climates (12). Lepidium
sensu stricto (excluding Cardaria Desv., Coronopus Zinn and Strogonovia
Kar. & Kir. includes 13 species and 2 subspecies that one of them L. vir-
ginicum is naturalized alien species in Turkey (6, 7). Cardaria, Corono-
pus and Stroganowia each include two taxa in Turkey. L. coronopus (L.)
Al-Shehbaz and L. didymium L. (formerly Coronopus), L. draba L. and
L. chalepense L. (formerly Cardaria), Stroganowia leventii V.I. Dorofeev
and S. leventii V.I. Dorofeev var. pilosa V.I. Dorofeev (new names of these
taxa are unsolved) (6, 13, 14, 15). In 1906, Thellung (16) defined the genus
Cardaria as a section of Lepidium, later however; Cardaria was accepted
as its own genus based on its indehiscent fruits (17, 12). Mummenhoff (18)
suggested that Cardaria should be classified within Lepidium according to
analyses of DNA sequence data. Other molecular research suggested that
not only Cardaria but also Coronopus, Stroganowia, Winklera Regel, and
Stubendorffia Schrenk ex Fisch., C.A. Mey. & Avé-Lall. are polyphyletic
and should be classified within Lepidium (9, 10, 19, 20, 21).
 M. BONA 33

Karyological studies on genus Lepidium show basic chromosome numbers

are x = 8 and chromosome number 2n = 16, 24, 28, 32, 40, 48, 64, 80 (22).
In this study detailed descriptions and identification key of the species
of Lepidium sensu stricto are given. Accepted names, type specimens and
synonyms of Lepidium updated according to current publications. Geo-
graphical distribution, habitat, specimen examined, and photographs are
also provided.
Materials and methods
Flowering and fruiting material of the genus Lepidium were collected
from different parts of Turkey for My PhD thesis (23) during May-August
2008, 2009 and 2010. Specimens were collected from as many different parts
of distribution area of the genus as possible in order to thoroughly examine
variation patterns. The specimens collected were kept at the Istanbul Univer-
sity, Department of Pharmaceutical Botany Herbarium (ISTE) and compared
with collections of ANK, E, GAZI, HUB, ISTF, ISTE, K, and VAN herbaria.
A range of characteristics of sepal, petal, stamen, silicula, pedicel,
septum, stigma and sinus that are considered to be taxonomically impor-
tant in the genus were investigated. For these investigations, all parts of
the specimens photographed by using MOTIC 2000 camera stereo micro-
scope system, and then measured by using Motic Image Plus 2.0 program.
These measurements were used for descriptions.
Results and disscussion
Lepidium L.
Annual, biennial or perennial herbs, rarely shrubs or lianas, glabrous,
papillose or with simple hairs. Stems erect, ascending or prostrate. Basal
leaves petiolate, simple, rosulate or not; cauline leaves short petiolate, ses-
sile, auriculate or amplexicaul. Sepals not saccate at base, oblong or ovate
(rarely orbicular), glabrous or pubescent. Petals oblanceolate, spathulate
or obovate, white or yellow, sometimes rudimentary or absent. Stamens 6,
4 or 2, filaments slender, neither winged nor toothed. Median nectar glands
present. Ovary with 2 (or 4) ovules; style short or absent. Fruits strongly
compressed a right angle to the septum or rarely inflated, dehiscent or rare-
ly indehiscent, elliptic, oblong, ovate or orbicular, winged or not, glabrous,
34 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

or pubescent; septum narrow broad, membranous, complete; style short or

long, exerted or included in apical notch or fruit not notched. Seeds pendu-
lous from apex of each loculus, winged or not, oblong or ovate, mucilagi-
nous, brown, black or green, radicle incumbent or obliquely so.
Dichotomous key for studied taxa
1. Fruits with a sinus at apex
2. Fruiting pedicel adpressed
3. Fruits 2–horned at apex 2. L. spinosum
3. Fruits emarginated without horn 3. L. sativum
2. Fruiting pedicel spreading
4. Cauline leaves amplexicaul
5. Cauline leaves ovate–orbicular,
flowers yellow 6. L. perfoliatum
5. Cauline leaves oblong–lanceolate, flowers
white 1. L. campestre
4. Cauline leaves not amplexicaul
6. Fruits ovate, petals absent 4. L. ruderale
6. Fruits orbicular, white petals present 5. L. virginicum
1. Fruits without a sinus
7. Stems with swellings at nodes, cauline leaves
1–2 pinnatisect 7. L. vesicarium
7. Stems without swellings at nodes, cauline leaves
not 1–2 pinnatisect
8. Leaves fleshy, basal leaves prominent at fruiting
time, fruit length more than 3 mm
9. Leaves auriculate, broadly ovate 10. L. cartilagineum
9. Leaves not auriculate, lanceolate or linear
10. Leaves subcylindrical, all linear 8. L. caespitosum
10. Leaves lanceolate, broad 9. L. pumilum
8. Leaves not fleshy, basal leaves mostly withered
at fruiting time, fruits less than 3 mm
11. Fruits pubescent, leaves laceolate-ovate,
auriculate 11. L. latifolium
11. Fruits glabrous, leaves oblong to linear, not auricılate
12. Fruits up to 2 mm, leaves glabrous 12. L. lyratum
12. Fruits 2–3 mm, leaves pubescent 13. L. graminifolium
 M. BONA 35

Description of the species

1. Lepidium campestre (L.) W.T. Aiton in Hort. Kew. ed. 2, 4: 88 (1812).
Synonym: Thlaspi campestre L. in Sp. PI. 646
Type: “Habitat in Europae arvis, viis argillosis, apricis” Herbarium Lin-
naeus No: 825:8 (lectotype designated by Al-Shehbaz (24)).
Annual or biennial, 15–60 cm, single or many stemmed, simple or much
branched, entire plant covered with short soft hairs. Basal leaves usually
lyrate, oblanceolate or ovate, 3.5–15 × 0.7–2.5 cm with 1–6.5 cm petiole.
Cauline leaves oblong-lanceolate, amplexicaul, sagitate at base. Upper
leaves smaller, simple, sessile, all leaves covered with short soft hairs.
Sepals white margined, 1.3–2 × 0.6–1.3 mm, purple on tip with short soft
hairs. Petals white, 1.7–3 × 0.4–1.1 mm. Stamens 6, 1.5–2.5 mm. Fruiting
pedicels erect, 3.5–6.3 mm with short soft hairs. Fruits covered with small
scale-like vesicles, strongly winged oblong, oblong to ovate, 5.5–7.5 ×
4–6.5 mm with 3.6–5 × 1.2–1.8 mm septum, emarginate at apex, sinus
0.5–1 mm wide; style exerted apical notch, stigma 0.4–0.7 × 0.2–0.25 mm.
Seeds wingless, ovate, 2.2–2.5 × 1.2–1.6 mm, blackish brown. Flowering
time Apr-Jun and fruiting time May-Jul (Figure 1).
Distribution: Europe, Caucasus; naturalized Australia and North and
South America.
Habitats: Field margin, roadside, around Quercus ssp. and Pinus ssp.
forests, s.l.–1700 m.
Chromosome number: 2n = 16 (22)
Specimen examined: A1 Edirne: Keşan, between Keşan –Enez, Kılıçköy, 20 m,
18.vii.2009, M. Bona 1118, ISTE 93396. A2(E) İstanbul: Sarıyer, between Rumeli Ka-
vağı–Rumeli Feneri, 100 m, 02.v.2010 M. Bona 1041, ISTE 93343. ibid., 27.v.2008, M.
Bona 1043, ISTE 93345. ibid., 09.v.2009, N 41 11 50 E 29 03 45, M. Bona 1087, ISTE
93367. ibid. 23.iv.2010, M. Bona 1122, ISTE 93401. A2(A) İstanbul: Yakacık, Aydos,
300 m, 25.iv.2008, M. Bona 1037, ISTE 93339. İstanbul: Yakacık, hills of Aydos Mt., 300
m, 25.iv.2008, M. Bona 1038, ISTE 93340. ibid., 10.v.2008, M. Bona 1042, ISTE 93344.
A5 Samsun: Lâdik, 900 m, fallow fields, 19.v.1964, C. Tobey, No: 589, E00372236. A7
Gümüşhane: prope Zigana in Ziganadagh, 20.vii.1889, Sintenis No: 1371. B5 Yozgat:
Akdağmadeni to Büyük Nalbant Dağ, at Yukarı Çuhalı, 2100 m, Beside irrigation ditch,
Flowers white. 6.vi.1965, Coode & Jones, No: 2067, E00372235.
36 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

Lepidium campestre is a unique taxon. It is easy to separate from other

Lepidium taxa because of its densely hairy stems and oblong-lanceolate,
amplexicaule leaves. Molecular studies showed that L. campestre related
with L. draba and L. appelianum (formerly Cardaria draba and C. pubes-
cens) (25, 26). Herbarium specimens also show sometimes L. campestre
could be missidentificated as L. draba. There are some diagnostic charac-
teristics to separate L. campestre from L. draba: L. campestre is annual with
long racemes, dehiscent and more or less oblong fruits, whereas L. draba is
perennial with corymbose panicle, indehiscent and ovate to cordate fruits.

Figure 1. Lepidium campestre: a- habit, b- infloresence.

2. Lepidium spinosum Ard. in Animad. Specim. In Alt. 2: 34, t. 16

(1764).
Synonym: L. cornutum Sibth. & Sm. in Prodr. Fl. Graec. 2: 6 (1813).
Type: not designated.
 M. BONA 37

Annual, 10–60 cm, single stemmed, branched from middle rarely up-
per part. Stem glabrous and waxy. Basal leaves 2-pinnatisect, 5–18 × 2–6
cm, with 1–5 cm petiole. Cauline leaves sessile, without auricles, linear to
lanceolate, slightly pinnate, upper linear and simple; all leaves with short
scattered hairs. Sepals white margined 1.3–1.9 × 0.6–1 mm, purplish on
middle–upper part, glabrous. Petals white, 2–3 × 0.7–1.4 mm. Stamens 6,
1.5–2.2 mm. Fruiting pedicels adpressed, 1.7–3 mm, glabrous. Fruits with
scattered hairs or glabrous, strongly winged oblong, oblong–ovate, 5–6 ×
2.2–2.3 mm, with 3–4 × 0.4–0.7 mm septum, deeply emarginate at apex,
sinus 0.8–1.8 mm wide; style included in sinus, stigma 0.3–0.5 × 0.2–0.3
mm. Seeds winged, ovate, 2–2.5 × 1–1.4 mm, brown. Flowering time Apr-
Jun and fruiting time Apr-Jun (Figure 2).
Distribution: Balkans, West Syria.
Habitats: Moist slopes, meadows, field margins, s.l.–1300 m.
Chromosome number: 2n = 16, 24 (22).
Specimen examined: A2(A) İstanbul: Kayışdağı, road of Küçükbakkalköy, Kayışdağı
Suyu, 12.v.1963, H. Demiriz, E00372544. B1 Balıkesir: Ayvalık, Alibey (Cunda) Island,
s.l., 24.iv.2009, M. Bona 1076, ISTE 93362. İzmir: Bergama, Asclepion, meadows, 1130
m 25.iv.2009, M. Bona 1078, ISTE 93363. C1 Muğla: Bodrum, 100 m, 11.iv.1965, Da-
vis 40905, E00372545. Muğla: Bodrum, between Ortakent–Bağla, roadsides, 170 m,
28.iv.2009, M. Bona 1079, ISTE 93364. Muğla: Bodrum, between Bodrum–Gümbet,
meadows, 40 m, 28.iv.2009, M. Bona 1080, ISTE 93365. C2 Marmaris: Cumali to Cni-
dus, 50–100 m, field weed, 17.iv.1965, Davis 41267, E00372546. Yatağan, Çine, 100 m,
granite hills, stem very tough, flowers white, 01.iv.1956, Davis & O. Polunin, No: 25562,
E00372559. C3 Antalya: Kumköy, between Antalya & Serik, 20 m, maquis on sandy soil,
flowers white, 06.v.1956, Davis & O. Polunin, No: 25724, E00372558. C5 Kahraman-
maraş: Road, Andirin to Çokak, 1000 m, Marshy ground, 18.v.1965, Coode & Jones, No:
1180, E00372546. C6 Adana: Bahçe (N. Amanus), Dumanlı Dağ near Haruniye, 700 m,
metamorphic banks, flowers white, 19.iv.1957, Davis & Hedge, No: 26875, E00372548.
Gaziantep: West of Fevzipaşa, West of Gaziantep, hills beside water, few, local, 600 m,
flowers white, 22.iv.1967, A.R.Mitchell, E00372552. Gaziantep: Fevzipaşa, 1800 ft.
15.iv.1935, E. K. Ball No: 2126. Hatay: Yayladağı, Kışlak, Yazı area, field margin, 900
m, 03.v.2009, M. Bona 1081, ISTE 93366. ibid., 12.v.2010, M. Bona 1128, ISTE 93407.
Lepidium spinosum is related to L. sativum. The general habitus are simi-
lar to L. sativum subsp. spinescens. L. spinosum is distinguished by a unique
oblong fruits and horned at apex. L. sativum subsp. sativum, L. sativum sub-
sp. spinescens and L. spinosum were grouped together acoording to mole-
culer data (25, 26, 27). In both cpDNA and ITS trees, L. spinosum and L.
sativum is sister to each other with more than 90% bootstrap support (28).
38 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

Figure 2. Lepidium spinosum: a- habit, b- infloresence.

3. Lepidium sativum L. in Sp. PL 644 (1753).

Annual, 10–70 cm, single stemmed, branched from middle or upper
in region of inflorescence. Stem glabrous and waxy. Basal leaves pinna-
tisect, 2-pinnatisect, lyrat, obovate or obovate–linear, 4.5–24 × 2–6 cm
with 1–13.5 cm petiole. Cauline leaves sessile, without auricles, slightly
pinnate-linear, pinnatisect, linear, simple or pinnatisect with linear seg-
ments. All leaves glabrous above and short soft hairy below and margin.
Sepals white margined 1.3–2 × 0.6–1.2 mm, purplish on middle-upper part
with scattered hairs. Petals white or lilac 2.3–3 × 0.9–1.3 mm. Stamens 6,
1.7–2.7 mm. Fruiting pedicels adpressed, 1.9–4.1 mm, glabrous. Fruits
glabrous, ovate, 4.3–6 × 3–4.8 mm with 3–4.4 × 0.5–1.2 mm septum,
emarginate at apex, sinus 0.3–0.7 mm wide; style exerted or included in
apical notch, stigma 0.2–0.6 × 0.2–0.3 mm. Seeds winged, ovate, 2.4–2.9
× 0.9–1.5 mm, brown. Flowering time Apr-Jun and fruiting time May-Jun
(Figure 3-4).
 M. BONA 39

1. Axes of fruiting racemes not ending in a spiny point,

stamens 6 (4 + 2), basal leaves without long linear seg-
ments……………………………………………………....subsp. sativum
1. Axes of fruiting racemes ending in a spiny point, sta-
mens 6 (equal), basal leaves with long linear seg-
ments………………………………………………...…..subsp.spinescens

Lepidium sativum L. subsp. sativum

Type: Herbarium Linnaeus No. 824.11 (lectotype designated by Hedge
(6)).
Distribution: Cultivated throughout most of the world.
Habitats: Cultivated as a salad, waste moist places.
Chromosome number: n = 2n = 16, 24 (22).
Specimen examined: A2(A) İstanbul: Maltepe, Dragos, 10 m, 10.v.2008, M. Bona
1040, ISTE 93342. ibid., 10.v.2009, M. Bona 1088, ISTE 93368. ibid, 24.iv.2010, M. Bona
1123, ISTE 93402. C6 Hatay: Antakya, near St. Peter’s Church, 150–300 m, 27.iv.1957,
Davis & Hedge, No: 27235, E00372269. Hatay: Altınözü, Fatikli, cultivated, 12.v.2010,
M. Bona 1129, ISTE 93408. B6 Kahramanmaraş: Türkoğlu, Büyük İmalı village, road-
side, 07.vi.2008, M. Bona 1064, ISTE 93354. Kahramanmaraş: Bertiz, Başdervişler vil-
lage, cultivated, 1270 m, 07.vi.2008, M. Bona 1067, ISTE 93355. C2 Muğla: Marmaris,
Söğüt to Bozburun, sea level–50 m, stony limestones places, 15.iv.1965, Davis 41173,
E00372279. C3 Isparta: Sütçüler, Çandır Village, cultivated, 21.iii.2010, M. Bona 1121,
ISTE 93399.
40 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

Figure 3. Lepidium sativum subsp. sativum: a- habit, b- infloresence.

Lepidium sativum L. subsp. spinescens (DC.) Thell. in Vierteljahr.

Naturf. Ges. Zürich, 51: 161 (1906).
Synonyms: L. spinescens DC. in Syst. 2: 534 (1821). L. sativum L. var.
spinescens (DC.) Jafri, in Fl. W. Pakistan 55: 55 1973.
Type: De Candolle Herbarium (GDC) G00203922
Distribution: Egypt, Syrian Desert, Iran.
Habitats: Waste moist places, basalt stony places.
Chromosome number: Unknown.

Specimen examined: A7 Şanlıurfa: Siverek, east of Siverek, c. 800 m, 19.v.1957, Da-

vis &Hedge, No: 28292, E00372264. Between Karacadağ and Siverek, around Damözü,
slopy meadows, 1250 m, 08.vi.2008, M. Bona 1071, ISTE 93357. ibid., 09.v.2010, M.
Bona 1125, ISTE 93404. Between Urfa and Siverek, 1841, Kostchy No:84. C3 Lydia:
Smyrna, in colibus ad Göstepe, 07.v.1906, J. Bornmüller No: 9067. C8 Mardin: 4 km
from Idil to Cizre, 590 m, 13.v.1966, Davis 42935, E00372285.
 M. BONA 41

Figure 4. Lepidium sativum subsp. spinescens: a- habit, b- infloresence.

According to Hedge (6), Lepidium sativum subsp. sativum and L. sativum
subsp. spinescens are only recognizably different from the type in the fruit-
ing stage. Some specimens are intermediate between the two subspecies.
This study provides some characteristics that are useful for distinguishing
the subspecies before the fruiting stage. L. sativum subsp. spinescens has
equal stamens and also has pinnatisect leaves with long linear segments.
42 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

4. Lepidium ruderale L. in Sp. PI. 645 (1753).

Type: Herbarium Linnaeus No. 824.17 (lectotype designated by Jonsell
and Jarvis (29)).
Annual 10–30 cm, simple or many stemmed, well branched above.
Stem densely or sparsely covered with short hairs. Basal leaves 2-pinnati-
sect, 2.5–6 × 0.6–1 cm with 1–3 cm petiole. Cauline leaves sessile, middle
without auricles, simple or slightly pinnate, upper simple, oblong-linear.
All leaves covered with short scattered hairs. Sepals white margined 0.7–
1.2 × 0.3–0.5 mm, purplish on middle–upper part. Petals absent. Stamens
2, 0.9–1.4 mm. Fruiting pedicels erect 2.2–4.6 mm covered short scattered
or densely hair. Fruits glabrous, , scarcely winged, ovate, 2–3 × 1.6–2.6
mm with 1.5–2.5 × 0.3–0.5 mm septum, emarginate at apex, sinus 0.2–0.5
mm wide; style included in sinus, stigma 0.1–0.2 × 0.1–0.2 mm. Seeds
wingless, ovate, 1.3–1.7 × 0.7–1 mm, brown. Flowering time Apr-Jun and
fruiting time Apr-Jun (Figure 5).
Distribution: Europe, Southwest Asia.
Habitats: Roadsides, waste places, field margins, s.l.–1550 m.
Chromosome number: 2n = 16, 32 (22)
Specimen examined: A9 Kars: Tuzluca, (Aras valey), 1070 m, brought by villager–
said to be wild, annual, 25.v.1966, Davis, 43582, E00372296. B4 Ankara: Yenimahalle,
Varlık District, around Ankara Stream, waste places, 850 m, 21.v.2009, M. Bona 1097,
ISTE 93377. ibid., 16.v.2010, M. Bona 1132, ISTE 93411. B7 Diyarbakır: around Öğret-
menevi, 10.v.2010, M. Bona 1126, ISTE 90405. Kayseri: Yeşilpınar, Ovaçiftliği, 1100
m, 30.vi.2009, M. Bona 1113, ISTE 93392. B8 Muş: between Bulanık and Karaçoban,
Şatırlar, Kırmızı Tuzla, 1600 m, 02.vi.2008, M. Bona 1044, ISTE 93346. B9 Bitlis:
1550 m, Castel ruins, 20.v.1966, Davis, 43368, E00372297. C6 Kahramanmaraş: Bertiz,
Başdervişler, 1300 m, 07.vi.2008, M. Bona 1068, ISTE 93356.
 M. BONA 43

Figure 5. Lepidium ruderale: a- habit, b- infloresence.

Lepidium ruderale is the only apetalous species in Turkey. It is widely

distributed but not collected much. L. ruderale is sister to L. pinnatifidum
in cpDNA (28) and other molecular researchs also shows they are related
(25, 26, 27). Their status in Turkey and relationship is discussed below.
5. Lepidium virginicum L. in Sp. PI. 645 (1753).
Synonym: L. iberis L. in Sp. PI. 645 (1753).
Type: Herbarium Linneaus No. 824.18 (lectotype designated by Marais
(30)).
Annual or biennial, 10–70 cm, single stemmed, branched above. Stem
covered with short soft hairs. Basal leaves pinnatisect, 2–13 × 1–4 cm,
petiole 1–6 cm, with short soft hairs. Cauline leaves without auricles, line-
ar–oblanceolate, sharply serrate, sparsely pubescent or glabrescent. Sepals
white margined 1–1.3 × 0.5–1 mm, sparsely pubescent, yellow–green on
middle–upper part. Petals white 1.5–2.1 × 1–1.5. Stamens 2(–3–4), 1.2–1.7
44 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

mm. pedicels erect, 2.2–4.5 mm, pubescent. Siliculae orbicular, winged,

2.5–3.6 × 2.4–3.5 mm, with 1.9–2.7 × 0.3–0.6 mm septum, emarginate at
apex, sinus 0.2–0.5 mm wide; style not exceeding to sinus, stigma 0.2–0.3
× 0.1–0.2 mm. Seeds strongly winged, orbicular 1.7–2 × 1–1.3 mm, brown.
Flowering time Mar–Jun and fruiting time May-Jul (Figure 6).
Distribution: Europe, North America, Australia.
Habitats: Roadsides, s.l.–350 m.
Chromosome number: 2n = 16, 32 (22)
Specimen examined: A7 Giresun: Tirebolu, 31.07.2008, M. Bona 1074, ISTE 93360.
ibid., 12.iii.2009, M. Bona 1101, ISTE 93381. Giresun: Tirebolu, between Gölağzı–Es-
piye, s.l., 18.v.2009, M. Bona 1091, ISTE 93371. Trabzon: Araklı, s.l., 16.v.2009, M.
Bona 1089, ISTE 93369. A8 Rize: The coastal path, s.l., 17.v.2009, M. Bona 1090, ISTE
93370. Rize: between Ardeşen–Pazar, 3 m, 22.vi.1981, A. Güner No: 3800, E00372388.
A9 Artvin: road of Artvin–Yusufeli, 45 km to Yusufeli, roadside, 350 m, 06.vi.2009, M.
Bona 1103, ISTE 93383.

Figure 6. Lepidium virginicum: a- habit, b- infloresence.

 M. BONA 45

Lepidium virginicum is an invasive alien plant in Turkey. It is native of

North America; introduced into South America, Europe, Asia, and Aus-
tralia. Herbarium specimens show it is distinguished with L. ruderale. L.
virginicum differs with small white flowers and orbicular fruits.
6. Lepidium perfoliatum L. in Sp. PI. 643 (1753).
Type: Herbarium Clifford 331, Lepidium 3 (lectotype designated by
Jonsell and Jarvis (28))
Annual or biennial, 10–35 cm, single stemmed, well branched above.
Stem sparsely pubescent lower and glabrous middle to upper part. Ba-
sal leaves 2–3-pinnatisect, pubescent, 4–15 × 1–3 cm, petiole 0.5–3.5 cm.
Cauline leaves simple, sessile, ovate–orbicular, perfoliate and glabrous.
Sepals yellowish margined 1–1.4 × 0.7–1 mm, sparsely pubescent, pur-
plish on middle-upper part. Petals yellow 1.2–1.8 × 0.3–0.5 mm. Stamens
6, 1–1.6 mm. Fruiting pedicel erect, 3–7 mm, glabrous. Fruits glabrous,
ovate, ovate-orbicular, winged, 3–6 × 2.7–5 mm, with 3–4.4 × 0.7–1.2 mm
septum, emarginate at apex, sinus 0.3–0.6 mm; style exerted apical notch,
stigma 0.2–0.4 × 0.1–0.2 mm. Seeds strongly winged, orbicular or ovate,
1.9–2.8 × 1.2–2 mm, brown. Flowering time Apr-Jun and fruiting time
Apr-Jul (Figure 7).
Distribution: Central and South Europe, Northwest and Central Asia,
South and West Australia.
Habitats: Field margins, waste places, roadsides, salty grounds, stony
slopes, s.l. 1850 m.
Chromosome number: n = 8; 2n = 16 (22).
Specimen examined: A4 Ankara: Road of Kızılcahamam–Güdül, 830 m, 13.iv.2008,
M. Bona 1019, ISTE 93322. Ankara: Güdül, roadsides, 740 m, 13.iv.2008, M. Bona 1021,
ISTE 93324. A5 Kırıkkale: Delice, between Delice and Sungurlu, 2 km to Sarıkaya Vil-
lage, salty places, 740 m, 19.v.2009, M. Bona 1093, ISTE 93373. Sinop: after Hacihamza
köy, 400 m, 11.v.1969, Tobey, No: 2610, E0001364. A7 Gümüşhane: Gümüşhane, stony
slopes, 1400 m, Stainton, No: 8326, E00051337. A8 Artvin: Yusufeli, 600 m, 14.vi.2010,
M. Bona 1140, ISTE 93416. B3 Afyonkarahisar: Road of Eskişehir–Afyon, 1000 m,
15.iv.2008, M. Bona 1027, ISTE 93330. Afyonkarahisar: Road of Afyon–İzmir, roadsides,
1000 m, 15.iv.2008, M. Bona 1028, ISTE 93331. Afyonkarahisar: Road of Afyon–İzmir
yolu, salty palces, 1000 m, 22.v.2009, M. Bona 1099, ISTE 93379. Eskişehir: Odun-
pazarı, Kozlubel, 940 m, 14.iv.2008, M. Bona 1023, ISTE 93326. Eskişehir: Sivrihisar,
1100 m, 14.iv.2008, M. Bona 1024, ISTE 93327. Eskişehir: Ertuğrul, 1000 m, 15.iv.2008,
46 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

M. Bona 1025, ISTE 93328. Eskişehir: Road of Eskişehir–Afyon, 85 km to Afyon, road-

sides, 15.iv.2008, M. Bona 1026, ISTE 93329. B4 Ankara: Şereflikoçhisar, edge of Tuz
Gölü, 900 m, 19.v.2009, M. Bona 1096, ISTE 93376. Ankara: Yenimahalle, around Anka-
mall, 840 m, 16.v.2010, M. Bona 1131, ISTE 93410. Konya: Kıreli, 1150 m, 18.iv.2008,
M. Bona 1034, ISTE 93336. B5 Kayseri: Yeşilpınar, Ovaçiftliği, 1080 m, 30.vi.2009,
M. Bona 1112, ISTE 93391. Kırşehir: Mucur, Seyfe Lake, 1100 m, 19.v.2009, M. Bona
1094, ISTE 93374. Yozgat: Himmetdede to Boğazlıyan, 20 km N of Himmetdede, 1200
m, marshy ground, probably saline, yellow flowers, beside stream, 25.v.1965, Coode &
Jones, No: 1481, E00051358. B6 Kahramanmaraş: Elbistan, c. 1100 m, on mud roofs, an-
nual, flowers white, 7.v.1957, Davis & Hedge, No: 27638, E00051340. B7 Elazığ: Elazığ
to Kale, (on Euphrates), 22 miles E. of Elazığ, 1300 m, Marly vineyards, 04.vi.1957.
Davis, No: 28934, E00051339. B8 Erzurum: city centre, roadsides, 1900 m, 14.vi.2010,
M. Bona 1138, ISTE 93415. Erzurum: Erzurum, Waste ground, 1900 m, 12.vi.1957,
Davis, No: 29245, E00051338. B9 Bitlis: Castel ruins, 1550 m, 20.v.1966, Davis, No:
43369, E00051328. Muş: between Bulanık–Karaçoban, Şatırlar, Kırmızı Tuzla, 1600 m,
02.vi.2008, M. Bona 1045, ISTE 93347. Van: Gürpınar, Çavuştepe, 1870 m, 03.vi.2008,
M. Bona 1057, ISTE 93350. Van: near the Van Lake, 03.vi.2008, M. Bona 1059, ISTE
93351. B10 Ağrı: 3 km N of Doğubeyazıt, 1700 m, meadows, 30.v.1966, Davis, No:
43853, E00051333. C3 Isparta: Eğridir, around Eğridir Lake, 1000 m, 15.iv.2008, M.
Bona 1029, ISTE 93332. Isparta: Şarkikaraağaç, Kızıldağ National Park, roadsides, 1150
m, 18.iv.2008, M. Bona 1032, ISTE 93334. Isparta: Şarkikaraağaç, 1100 m, 18.iv.2008,
M. Bona 1033, ISTE 93335. C4 Karaman: Road of Karaman–Karapınar, Karapınar, 1000
m, 19.iv.2008, M. Bona 1035, ISTE 93337. C5 İçel: Tarsus, Çukurova, Çakırlı village,
9.iv.1967, Chester F. Deaver, T 99, E00051343.

Lepidium perfoliatum is only species which has yellow flowers. Be-

cause of that it is easy to seperate from other Turkish Lepidium taxa. Her-
barium specimens showed L. perfoliatum confused with Thlaspi perfolia-
tum L. L. perfoliaum has ovate or ovate–orbicular silicula and one seed
in each loculus. Acording to molecular research L. perfoliatum is related
with L. draba, and L. campestre (25; 28). All these Lepidium taxa are char-
acterized by auriculate and amplexicaul cauline leaves, flowers with 2 +
4 stamens, and all taxa are native in the Mediterranean area, Europe, and
southwest and central Asia (25).
 M. BONA 47

Figure 7. Lepidium perfoliatum: a- habit, b- infloresence.

7. Lepidium vesicarium L. in Sp. PI. 643 (1753).

Lectotype: Ic. “Lepidium orientale, Nasturtii folio, caule vesicario” in
Buxbaum (1728: 26) (31) (lectotype designated by Lopez Gonzalez (32)).
Annual or biennial, 15–70 cm, single stemmed, branched in region of
inflorescence. Stems with prominent turbinate swellings at nodes, gla-
brous, waxy. Basal leaves 2–3 pinnatisect with linear segments, 5–12 ×
2–5 cm, petiole 1–2.5 cm. Cauline leaves sessile, without auricles, 1–2 pin-
natisect with linear segments. All leaves glabrous above pubescent below.
Sepals white margined 1–1.4 × 0.7–1 mm, yellow-green on middle-upper
part, scarcely pubescent. Petals white 2.2–2.9 × 1.3–1.6 mm. Stamens 6,
1.4–1.7 mm. Fruiting pedicel erect, 4–6.5 mm, villous, glabrescent at fruit-
ing time. Fruits glabrous, ovate–elliptic, scarcely winged at apex, 2.5–3.5
× 2.2–2.8 mm, with 2–3 × 0.7–1.1 mm septum, without sinus, slightly
notched, stigma 0.4–0.5 × 0.4–0.5 mm. Seeds wingless, oblong–elliptic,
1.8–2.2 × 1–1.3 mm, green. Flowering time May–Jun and fruiting time
Jun-Jul (Figure 8).
48 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

Distribution: Caucasus, Iran.

Habitats: Stony slopes, roadsides, 800–1900 m.
Chromosome number: 2n = 16, 32 (22)
Specimen examined: A8 Erzurum: Yusufeli–Erzurum road, Artemisia steppe,
09.vii.1960, Stainton & Henderson, No: 6107, E00051392. 12 miles W. of Erzurum,
5000 ft. vii.1965, E.M. Rix No:24. A9 Kars: Kağızman to Akçay (Aras valley), 1100–
1200 m, 19.vii.1966, E00051283. B9 Ağrı: Ağrı to Doğubeyazıt, 44 E, 39 8 N, 1750
m, 17.vii.1967, David Valton, E0051389. Ağrı: Doğubeyazıt, Suluçem Village, 1730 m,
16.viii.2009, M. Bona 1120, ISTE 93398. Iğdır: Between Iğdır & Doğubeyazıt, 2 miles
from Iğdır, 11.viii.1970, Hewitt No: 338, E00051376. Van: Havasor–Hoşap, roadsides,
30.vii.1954, Davis & O. Polunin, No: 23284a, E00051393. Van: Gürpınar, Çavuştepe,
1870 m, 03.vi.2008, M. Bona 1055, ISTE 93349. ibid., 18.v.2010, M. Bona 1133, ISTE
93412. Van: road to Üçler Village, 19.v.2010, M. Bona 1137, ISTE 93414.

Lepidium vesicarium has unique character because of the distinctive

nodal swelings. It is distributed only eastern part of Turkey and has no
close relatives.

Figure 8. Lepidium vesicarium: a- habit, b- infloresence.

 M. BONA 49

8. Lepidium caespitosum Desv. in J. Bot. Appl. 3: 165 & 178 (1814).

Synonym: L. cartilagineum (J. May.) Thell. subsp. caespitosum (Desv.)
Thell. In Die Gattung Lepidium, op. cit., 156 (1907).
Type: in Paludibus Armeniae prope Erzerum, Tournefort ! (type desig-
nated by Hedge (6)).
Perennial, 8–20 cm, many stemmed, branched above. Petiolar remains
present at base of stem, underground part often swollen. Lover part of
stem glabrous, papillate in region of inflorescences or all stem sparsely
pubescent. Basal leaves subcylindrical, linear, 3–8 × 0.1–0.3 cm, petiole
1–2.5 cm. Cauline leaves sessile, without auricles subcylindrical, linear.
All leaves glabrous. Sepals white margined 1.5–3 × 1–2 mm, glabrous,
purplish on upper part. Petals white 2.5–5 × 1.5–2.5 mm. Stamens 6, 2–3
mm. Fruiting pedicel erect, 4–7.4 mm, sparsely pubescent or papillate.
Fruits glabrous ovate, scarcely winged 2.7–4.5 × 2.3–3.8 mm, with 2.5–
3.5 × 0.6–1 mm septum, without sinus, stigma 0.3–0.5 × 0.2–0.3 mm.
seeds scarcely winged, ovate, 1.8–2.2 × 1–1.4 mm, brown. Flowering time
May-Jun and fruiting time Jun-July (Figure 9).
Distribution: Endemic.
Habitats: Salty marshes, 900–1800 m.
Chromosome number: Unknown.
Specimen examined: A8 Erzurum: road of Artvin, marshy palces, 1750 m, 07.vi.2009,
M. Bona 1106, ISTE 93385. B4 Ankara: Şereflikoçhisar, edge of Tuz Gölü, 1000 m,
20.vii.2008, M. Bona 1072, ISTE 93358 ibid., 19.v.2009, M. Bona 1095, ISTE 1095. Kon-
ya: Road of Ankara, between Ağabeyli and Kırkışla, salty marshes, 950 m, 23.v.2009, M.
Bona 1100, ISTE 93380. Konya–Cihanbeyli, 81 km, fringe of Ağabeyli Köyü, in the vi-
cinity of Aci Tuz Gölü (Bulok Gölü), 980 m, 27.v.1956, I.C. Hedge No: 2770, E00051318.
B5 Nevşehir: Sifegöl, nr. Mucur, 17.vi.1954, Davis No: 21795, E00051319. B6 Sivas:
Zara–Sivas road, 1300 m, 20.v.1960, Stainton & Henderson No: 5086, E00051322.

According to Hedge (1965), with the limited material collected it is

apparently a constant taxon, but Hedge (1965) suggested that further study
of more abundant material might show that it is not specifically different
from Lepidium cartilagineum. This study investigated more specimens
and shows L. caespitosum is clearly a different species.
50 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

Figure 9. Lepidium caespitosum: a- habit, b- infloresence.

9. Lepidium pumilum Boiss. & Bal. in Boiss., Diagn. ser. 2(6): 21
(1859).
Synonym: L. cartilagineum subsp. (J. May.) Thell. pumilum (Boiss. &
Bal.) Hedge in Fl. Iran 57: 67 (1965).
Type: Herbarium Kew, K000642709, K000642710 (Balansa 999)!
(type designated here).
Perennial, 8–17 cm, many stemmed, branched above. Petiolar remains
present at base of stem. Lover part of stem glabrous, upper part pubescent
or all stem sparsely pubescent. Basal leaves elliptic 3–9 × 0.3–1.5 cm, peti-
ole 1–3 cm, sparsely pubescent or glabrous. Cauline leaves sessile, without
auricles, linear–lanceolate. All leaves glabrous. Sepals white margined 1.5–
2 × 1–1.5 mm, sparsely pubescent, purplish on upper part. Petals white 2.5–
3.2 × 1.2–1.6 mm. stamens 6, 1.8–2.4 mm. Fruiting pedicel erect, 3.5–6.5
mm, pubescent. Fruits glabrous, ovate, scarcely winged, 2.7–3.7 × 2.4–3.4
mm, with 2.6–3 × 0.7–0.9 mm septum, without sinus, stigma 0.4–0.7 ×
0.2–0.4 mm. Seeds scarcely winged, ovate, 1.9–2.1 × 1.2–1.4 mm brown.
Flowering time May-Jun and fruiting time Jun-Jul (Figure 10).
Distribution: Europe, South Russia.
Habitats: Salty damp places, 600–1200 m.
Chromosome number: 2n = 16 (22).
 M. BONA 51

Specimen examined: A5 Çorum: İskilip, 26 miles from Çorum to İskilip, Kızılırmak

valley, saline marshy ground, 600 m, 29.v.1965, Coode & Jones No: 1725, E00051316.
Kırıkkale: Delice, between Delice and Sungurlu, salty grounds, 750 m, 19.v.2009, M.
Bona 1092, ISTE 93372. B4 Ankara: 10–12 km NE of Koçhisar, edge of Tuz Gölü, 900
m, 29.iv.1966, Davis No: 42175, E00051315. B5 Kayseri: between İncesu and Yahyalı,
salty grounds, 1070 m, 30.vi.2009, M. Bona 1109, ISTE 93388. Kayseri: Tuz Gölü, 1150
m, 30.vi.2009, M. Bona 1115, ISTE 93400. Yozgat: Himmetdede to Boğazlıyan, 20 km N
of Himmetdede, 1200 m, 25.v.1965, Coode & Jones, No: 1470, E00051314. C5 Niğde:
Cappadocia, 1100 m, 03.vi.1898, W. Siehe No: 46.

Figure 10. Lepidium pumilum: a- habit, b- infloresence.

10. Lepidium cartilagineum (J. May.) Thell. in Vierteljahr. Naturf. Ges.

Zürich 51: 173 (1906).
Synonym: Thlaspi cartilagineum J. May. in Abhandl. Böhm. Ges., 235.
t. 7 (1786). Lepidium crassifolium Waldst. & Kit. In PL Rar. Hung. 1: 4,
t. 4 (1799). Lepidium cartilagineum (J. May.) Thell. subsp. crassifolium
52 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

(Waldst. & Kit.) Thellung in Neue Denkschr. Schw. Ges. Naturv. 41: 153
(1906); L. crenatifolium Boiss. & Bal. in Boiss., Diagn. ser. 2(6): 22 (1859).
Type: Thlaspi cartilagineum; capsulis subrotundis integris, foliis ova-
tolanceolatis cartilagineis, [fl., fr. prim.] / Ex herbario Meyeriano” (PR)
(type Designated by German (32)).
Perennial, 10–35 cm, ascending, many stemmed, branched above. Peti-
olar remains present at base of stem. Lower part of stem glabrescent, upper
part pubescent, rarely all stem glabrecent. Basal leaves orbicular, ovate, ±
acuminate, sometimes scarcely crenulate, rarely almost pinnatifid, 3.5–10
(–25) × 1–4 (–14) cm. Petiole 1.5–4.5 (–8.5) cm, sparsely pubescent. Caul-
ine leaves sessile, auriculate, broadly ovate or elliptic. All leaves glabrous.
Sepals white margined 1.5–1.9 × 0.9–1.3 mm, purplish on middle–upper
part, scarcely pubescent. Petals white 2–3 × 1–2 mm. Stamens 6, 1.4–2
mm. Fruiting pedicel erect, 3.5–6.5 mm, pubescent rarely glabrous. Fruits
glabrous, ovate, scarcely winged, 2.3–3.7 × 2–3.2 mm, with 1.9–3.3 ×
0.5–0.8 mm septum, without sinus, stigma 0.3–0.6 × 0.2–0.5 mm. Seeds
scarcely winged, ovate, 2–2.4 × 1.2–1.5 mm, brown. Flowering time May-
Jun and fruiting time Jun-Jul (Figure 11).
Distribution: Europe, South Russia, Caucasus, Iran, Afghanistan, Cen-
tral Asia.
Habitats: Salty grounds, 800–1800 m.
Chromosome number: 2n = 16, 28, 40 (22).
Specimen examined: A8 Erzurum: between Erzurum–Ilıca, salty places, 1750 m,
07.vi.2009, M. Bona 1107, ISTE 93386. Circa Erzeroum, vi.1853, Huet du Pavillon. A9
Kars: 3 km E of Iğdır DÜÇ (Aras valley), 800 m, salty flats, perennial, flowers white,
29.v.1966, Davis No: 43838, E00051308. Aralık, (Aras valley), 810 m, 26.v.1966, Davis
No: 43643, E00051299. B3 Afyonkarahisar: road of Afyon–İzmir, salty places, 1000 m,
22.v.2009, M. Bona 1098, ISTE 93378. B4 Niğde: Tuz Gölü, N.W. of Ulukışla, (N.W.
of Aksaray), 900–950 m, salt steppe, perennials, fruits angustiseptate, 2 seeded, leaves
fleshy, 01.ix.1957, Davis & Hedge No: 32830, E00051310. B5 Kayseri: between İncesu–
Yahyalı, salty places, 1070 m, 30.vi.2009, M. Bona 1110, ISTE 93389. Kayseri: Yeşilpınar,
Ovaçiftliği Village, 1070 m, 30.vi.2009, M. Bona 1114, ISTE 93393. Yozgat: Himmetd-
ede to Boğazlıyan, 20 km N of Himmetdede, 1200 m, marshy ground, probably saline,
25.v.1965, Coode & Jones, No: 1474, E00051305. B9 Ağrı: 3 km N of Doğubeyazıt,
damp, salty flats, 1700 m, 30.v.1966, Davis No: 43851, E00051297. Van: Van Y.Y. Uni-
versity, 1650 m, 04.vi.2008, M. Bona 1061, ISTE 93352. ibid. 18.v.2010, M. Bona 1136,
ISTE 93413. Van: 4 km W of Erciş, 1720 m, 22.v.1966, Davis No: 43451, E00051296.
 M. BONA 53

Figure 11. Lepidium cartilagineum: a- habit, b- infloresence.

Lepidium pumilum and L. cartilagineum are represented in Flora of
Turkey as two subspecies of L. cartilagineum which are L. cartilagineum
subsp. cartilagineum and L. cartilagineum subsp. crassifolium (6). Later,
Hedge (33) proposed a new combination, L. cartilagineum subsp. pumilum
for the plants with narrow and non-auriculate leaves and L. cartilagineum
subsp. cartilagineum for the plants with wide auriculate leaves and placed
L. crassifolium into a synonymy of the L. cartilagineum subsp. cartilag-
ineum. In this study synonyms of taxa are given according to improved
version of the Flora Iranica (33). Because, according to Mayer (34) Thlas-
pi cartilagineum has wide and airuculate leaves. According to Thellung
(18) T. cartilagineum is synonym of L. cartilagineum subsp. crassifolium.
My observations on the herbarium specimens support that results. Wrong
typification and using synonims of these taxa still continiue; L. cartilag-
ineum subsp cartilagineum and L. cartilagineum subsp. crassifolium are
represented as two different taxa (35). Beside this, L. crenatifolium which
has ovate leaves (36) is represented as synonym of L. cartilagineum subsp.
crassifolium (35).
54 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

Herbarium specimens show that L. caespitosum, L. pumilum and L. car-

tilagineum are easily separated from each other, from stem length, leaf
shape, and auricle characteristics. These characteristics are useful for both
young and mature specimens. The taxa also have different habitats. L.
caespitosum grows in marsh places, L. pumilum grows damp places, and
L. cartilagineum grows mostly dry grounds. So these three taxa should be
considered as separated species.
11. Lepidium latifolium L. in Sp. PL 644 (1753).
Type: Herbarium Linneaus No. 824.11a (LINN) (lectotype designated
by Jafri (37)).
Perennial, 30–100 cm, stoloniferous, much branched in region of in-
floresence. Stem glabrous. Basal and lower leaves lanceolate, ovate,
toothed at margin 10–20 × 3.5–5.5 cm, pubescent, petiole 4–7 cm. Cauline
leaves sessile very rarely short petiolate, without auricles, lanceolate, sim-
ple or toothed at margin, glabrous. Upper leaves bract–like. Sepals white
margined 1–1.5 × 0.9–1 mm, purplish on middle–upper part, sparsely
pubescent. Petals white 2.3–2.7 × 0.9–1.2 mm. Stamens 6, 1.7–2.2 mm.
Fruiting pedicel erect, 3.5–5 mm, glabrous. Fruits pubescent, orbicular,
wingless, 1.8–2.2 × 1.6–2 mm, with 1.7 × 0.5 mm septum, without sinus,
stigma 0.15–0.2 × 0.2–0.3 mm. Seeds scarcely winged, ovate, 0.8–1 ×
0.5–0.7 mm, brown. Flowering time Apr-Aug and fruiting time Jul-Sep
(Figure 12).
Distribution: Europe, Southwest Asia, North Africa, Tasmania, South
Australia.
Habitats: Marshy palces, field margins, s.l. 2000.
Chromosome number: 2n = 24 (22).
Specimen examined: A5 Amasya: Galatia, Madissadjian No: 774. A9 Ağrı: Diyadin,
at hot springs, c. 6000 ft., 09.viii.1970, Hewitt No: 255, E00051266. Kars: 5–10 km E
of Iğdır DÜÇ (Aras valley), 800 m, edge of Phragnites marsh, perennials, sepals pur-
plish, 29.v.1966, Davis No: 43834, E00051264. B3 Kütahya: Eskişehir to Kütahya 23.
Km from K., 800–900 m, banks of river Porsuk, pale pink to white flowers, 22.vi.1962,
Dudley No:36008, E00372311. Gediz, 850 m, waste ground, perennial, flowers white,
04.iv.1962, Davis & Code No: 36904, E00051270. B4 Ankara: Çandır, E of main road to
Çankırı, 870 m, 40 13 52.0 N 33 27 36.5 E, 12.vii.1999, R.D. Reeves, A.R. Kruckeberg, N.
Adıgüzel No:1904, E00321655. Konya: Çumra road, 1000 m, 21.07.2009, M. Bona 1073,
ISTE 93359. B5 Kayseri: Yeşilpınar, Ovaçiftiliği, 1080 m, 30.vi.2009, M. Bona 1111,
 M. BONA 55

ISTE 93390. Kayseri–İncesu, 1200 m, marsh, perennial, 29.viii.1957, Davis & Hedge
No: 32754, E00051265. B5 Niğde: Nevşehir, 1200 m, damp hollow, petals white, calyx
pale purple, 21.vi.1952, Davis No: 19098, E00372307. B8 Erzurum: W. Of Tercan, 1500
m, Edge of marshy hollow, Perrenial, Flowers white, 11.vii.1957, Davis & Hedge No:
30946, E00051271. B9 Van: Sanayi district, near Morali stream, 1650 m, 04.vi.2008, M.
Bona 1063, ISTE 93353. C8 Mardin: in ruderatis, 09.vi.1888, P. Sintenis No: 1084. C9
Hakkari: 5 km from Şemdinli to Yüksekova, 1550 m, bank of stream, perennial, flower
buds white, 16.vi.1966, Davis No: 45020, E00372322. 12 km from Yüksekova, riverside
scree, 2000 m, 04.ix.1967, Duncan & Tait, No: 159, E00051253.

Figure 12. Lepidium latifolium: a- habit, b- infloresence.

12. Lepidium lyratum L. in Sp. PL 644 (1753).

Type: “Lepidium orientale, Nasturtii Crispi folio” in Tournefort (38)
(lectotype designated by Hedge (39)).
Perennial, 35–75 cm, many stemmed, much branched in region of inflore-
cense. Stem glabrous and waxy. Basal leaves lyrat with toothed margin, 6.5–
12 × 1.5–3.5 cm, petiole 2.5–5 cm. Cauline leaves sessile, without auricles,
oblong, linear, simple or toothed at apex. All leaves glabrous. Sepals white
56 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

margined 0.9–1.2 × 0.6–0.8 mm, purplish on middle-upper part, sparsely pu-

bescent. Petals white 1.7–2.5 × 1–1.5 mm. Stamens 6, 1.4–1.6 mm. Fruiting
pedicel erect 2.5–4 mm, glabrous or sometimes scarcely pubescent. Fruits
glabrous, orbicular, wingless 1.6–2 × 1.5–2 mm, with 1.5–1.6 × 0.4–0.6
mm septum, without sinus, stigma 0.2–0.3 × 0.2–0.3 mm. Seeds wingless or
scarcely winged, narrowed elliptical 1.3–1.4 × 0.6–0.7 mm, brown. Flower-
ing time May-Aug and fruiting time Jun-Aug (Figure 13).
Distribution: South Russia, Caucasia, Southwest Asia.
Habitats: Crevices of rocks, 400–1800 m.
Chromosome number: Unknown.
Specimen examined: A9 Artvin: Yusufeli, 600 m, 06.vi.2009, M. Bona 1104, ISTE
93384. ibid., 14.vi.2010, M. Bona 1141, ISTE 93417. Artvin: Artvin–Yusufeli road, 45
km to Yusufeli, roadsides, 450 m, 06.vi.2009, M Bona 1102, ISTE 93382. Erzurum: 14
km from Tortum to Oltu, 1500 m, Habit of Gypsophila, 29.vii.1966, Davis No: 47538,
E000372328. C9 Hakkari: Cilo Dag, in Diz deresi, 5700 ft. Gravel terrace. 06.viii.1954,
Davis & Polunin No: 23933.

Figure 13. Lepidium lyratum: a- habit, b- infloresence.

 M. BONA 57

Lepidium lyratum is similar with L. persicum Boiss. subsp. persicum,

and L. lacereum C. A. M., species that do not occur in Turkey. Some mor-
phological characters might be useful to separate these taxa from L. lyra-
tum. L. persicum subsp. persicum has almost leafless on the middle and
upper part of the stem, leaves densely pubescent, and basal leaves line-
ar-lanceolate. Stigma length is larger than width in L. lacereum. L. lyra-
tum group and L. persicum need detailed investigation with more matierial
which should include all Caucasus specimens.
13. Lepidium graminifolium L. Syst. Nat. ed. 10, 2: 1127 (1759).
Type: Herbarium Linneaus No. 824.15 (LINN) (lectotype designated
by Hedge (39).
Perennial, (–10)30–50(–80), many stemmed, much branched above.
Stem glabrous or rarely sparsely pubescent. Basal leaves lyrat, 3–8 × 0.5–
1.5 cm, petiole 1.5–2.5 mm. Cauline leaves sessile, without auricles, lin-
ear. All leaves pubescent. Sepals white margined 0.9–1.2 × 0.7–0.9 mm,
purplish on middle-upper part, scarcely pubescent. Petals white 1.2–1.6
× 0.7–0.9 mm. Stamens 6, 1.2–1.6 mm. Fruiting pedicel erect, 2.6–4.5
mm, glabrous or rarely scarcely pubescent. Fruits glabrous, ovate-elliptic,
scarcely winged, 2–2.8 × 1.7–2.3 mm, with 1.6–2.1 × 0.6–0.8 mm sep-
tum, without sinus. Stigma 0.2–0.3 × 0.2–0.3 mm. Seeds wingless, elliptic,
1.3–1.5 × 0.7–0.9 mm, brown. Flowering time Apr-Sep and fruiting time
Apr-Sep (Figure 14).
Distribution: Europe, Northwest Africa, Southwest Asia.
Habitats: Dry slopes, s.l.–1800 m.
Chromosome number: 2n = 16, 48 (22).
Specimen examined: A2(E) İstanbul: Kasımpaşa, roadsides, 16.vii.2009, M. Bona
1116, ISTE 93394. İstanbul: Küçükçekmece, s.l., 10.vi.2009, M. Bona 1108, ISTE 93387.
İstanbul: Şişli, Mecidiyeköy, Gülbağ, 17.vii.2009, M. Bona 1117, ISTE 93395. A2(A) İs-
tanbul: Maltepe, Dragos, 10 m, 24.iv.2009, M. Bona 1124, ISTE 93403. İzmit: 10 m, dry
S. slopes, 08.ix.1957, Davis & Hedge No: 32916, E00372385. A3 Zonguldak: Karabük,
300 m, garden weed, flowers white, 05.viii.1962, Davis & Code No: 38965, E00372383.
A6 Samsun: Road bank east of Samsun, 5 m, ix.1964, E00372382. B1 Çanakkale: Kilit-
bahir, 19.vii.2009, M. Bona 1119, ISTE 93397. Çanakkale: Dardanelles, 1883, P. Sintenis
No:566. C6 Hatay: Belen, Güzelyayla Village, 750 m, 20.viii.2009, M. Bona 1075, ISTE
93361. Hatay: Belen, Güzelyayla, serpentine area (Soğukoluk), 800 m, 36 29.15–36 N,
36 09.82–96, E, 05.viii.2001, R.D. Revees 2177, N. Adıgüzel & Ö. Bingöl (NA4052),
E00321664.
58 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

According to Lee (28) PI intron gene tree and the evolution of lateral
stamen arrangements show that Lepidium graminifolium and L. lyratum
closely related to each other. Herbarium specimens and morphological in-
vestigations also show that L. graminifolium is close to L. lyratum but it
differs from its much longer fruits and pubescent leaves.

Figure 14. Lepidium graminifolium: a- habit, b- infloresence.

Flowering time of the studied taxa is from March to September and
fruiting time is from April to September. Lepidium species distributed from
sea level to 2000 m altitude, except L. vesicarium, L. caespitosum, L. pum-
ilum, L. cartilagineum, and L. lyratum. Distribution of the taxa starts with
400 m at least. The studied taxa are mostly founded at waste places, field
margins, and roadsides. Three of them have special habitats; L. caespito-
sum, L. pumilum, and L. cartilagineum are grown only salty places.
Lepidium hirtum (L.) Sm. was recorded East Aegean Islands and it has
not been recorded from Turkey. L. pinnatifidum Ledeb. was published as a
new record (40) for Turkey. This new record was collected locality which
 M. BONA 59

gave in the publication and specimens of the researchers also investigat-

ed. Besides this L. pinnatifidum and L. ruderale specimens at Edinburgh
and Kew Herbaria were compared. All these researches showed that this
record was incorrect and collected material is L. ruderale. L. pinnatifidum
is easily recognized from L. ruderale by its small white petals and almost
glabrous stems.
As a conclusion; This study recognises 14 Lepidium s.s. taxa from Tur-
key. New characteristics are point outed for separate to L. sativum subsp.
sativum from L. sativum subsp. spinescens. Lepidium pumilum and L. car-
tilagineum are recognized at species rank. Type specimen is designated
for L. pumilum. This study also shows that L. pinnatifidum does not occur
in Turkey, although previously it had been considered a new record for
Flora of Turkey. Besides that, in this study detailed descriptions and iden-
tification key of the species of Lepidium sensu stricto are given. Accepted
names, type specimens and synonyms of Lepidium updated according to
current publications. Geographical distribution, habitat, specimen exam-
ined, and photographs are also provided.

Acknowledgements
I want to thank you Doç Dr. Aliye Aras for her support to finishing my PhD thesis.
Thanks are due to the curators and staff of all the herbaria listed under Material and Meth-
ods, especially E in Edinburgh and K in London. Authors also want to thank Sibbald Trust
who gave financial support to herbarium and library resaerchs in Edinburgh and London.
This study was supported by Istanbul University Research Fund (Project No: 2122).

References
1. Al-Shehbaz IA (2012). A generic and tribal synopsis of the Brassicaceae (Crucifer-
ae). Taxon 61: 931-954.
2. Koch MA, Mummenhoff K (2006). Editorial: Evolution and phylogeny of the Bras-
sicaceae. Plant Syst Evol 259: 81–83.
3. Candolle P (1821). Regni Vegetabilis Systema Naturale, Volume 2.
4. Schulz OE (1936). Cruciferae. In: Engler A, Harms H (eds.). Die Naturlichen Pflan-
zenfamilien ed. 2, 17B: 227-658. Leipzig: Verlag von Wilhelm Engelmann.
5. Beilstein MA, Al-Shehbaz IA & Kellogg EA (2006). Brassicaceae phylogeny and
trichome evolution. Am J Bot 93: 607-619.
60 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

6. Hedge I (1965). Lepidium L. – In: Davis PH (ed.), Flora of Turkey and the East Ae-
gean Islands Volume I. Edinburgh, UK: Edinburgh University Press.
7. Davis PH, Mill RR Tan K (1988). Flora of Turkey and the East Eagean Islands Vol-
ume X. Edinburgh Univ. Press.
8. Güner A, Özhatay N, Ekim T, Başer KHC (2000). Cruciferae. Flora of Turkey and the
East Aegean Island Volume XI. Edinburgh, UK: Edinburgh University Press.
9. Koch M, Al-Shehbaz IA, Mummenhoff K (2003). Molecular systematics, evolution,
and population biology in the mustard family (Brassicaceae). Ann Missouri Bot Gard
90: 151–171.
10. Al-Shehbaz IA, Beilstein MA, Kellogg EA (2006). Systematics and phylogeny of the
Brassicaceae (Cruciferae): an overview. Plant Syst Evol 259: 89–120.
11. Al-Shehbaz IA, Mutlu B, Dönmez AA (2007). The Brassiceae (Cruciferae) of Tur-
key, updated. Turk J Bot 31: 327–336.
12. Al-Shehbaz IA (1986). The genera of Lepidieae (Cruciferae; Brassicaceae) in the
southeastern United States. J Arnold Arbor 67: 265–311.
13. Dorofeev VI, Korotyaev BA, Gültekin L (2004). A new species of the genus Stro-
ganowia Kar. et Kir. (Cruciferae) from Northeast Turkey and Rhynchophorous bee-
tles (Coleoptera, Curculionoidea) associated with it. Byull Moskovsk Obshch Isp Prir
Otd Biol 109: 72-76.
14. Dorofeev VI (2006). Taxa nova in Stroganowia leventii V.I. Dorof. (Cruciferae). No-
vosti Sist Vyssh Rast 38: 127-130.
15. Yüzbaşıoğlu İS, Keskin M (2013). A new record for the flora of Turkey: Lepidium
didymum L. (Brassicaceae). Biodicon, 6, 46–48.
16. Thellung A (1906). Die Gattung Lepidium (L.) R. Br. Verlag von Georg & Co.
17. Mulligan GA, Frankton C (1962). Taxonomy of the genus Cardaria with particular
reference to the species introduced into North America. Can J Bot 4: 1411–1425.
18. Mummenhoff K (1995). Should Cardaria draba (L.) DESV. be classified within the
genus Lepidium L. (Brassicaceae)? Evidence from subunit polypeptide composition
of RUBISCO. Feddes Repert 106: 25–28.
19. Al-Shehbaz IA, Mummenhoff K, Appel O (2002). Cardaria, Coronopus, and Stro-
ganowia are united with Lepidium (Brassicaceae). Novon 12: 5–11.
20. Bailey CD, Koch MA, Mayer M, Mummenhoff K, O’Kane SL, Warwick SI, Wind-
ham MD Al-Shehbaz IA (2006). Towards a global nrDNA ITS phylogeny of the
Brassicaceae. Mol Biol Evol 23: 2142–2160.
21. Al-Shehbaz IA, Mummenhoff K (2011). Stubendorffia and Winklera belong to the
expanded Lepidium (Brassicaceae). E J B 68: 165–170.
 M. BONA 61

22. Warwick SI, Al-Shehbaz IA (2006). Brassicaceae: Chromosome number index and
database on CD–Rom. Plant Syst Evol 259: 237–248.
23. Bona M. (2010). Türkiye’nin Lepidıum L. (Tere) (Brassicaceae) Türleri Üzerinde
Morfolojik ve Taksonomik Araştırmalar. Doktora Tezi. İstanbul Üniversitesi, Fen
Bilimleri Enstitüsü.
24. Al-Shehbaz IA (2010). A synopsis of the South American Lepidium (Brassicaceae).
Darviniana 48: 141-167.
25. Mummenhoff K, Brüggemann H, Bowman JL (2001). Chloroplast DNA phylogeny
and biogeography of Lepidium (Brassicaceae). Am J Bot 88, 2051-2063.
26. Mummenhoff K, Polster A, Mühlhausen A, Theißen G (2009). Lepidium as a model
system for studying the evolution of fruit development in Brassicaceae. J Exp Bot
60, 1503-1513.
27. Mummenhoff K, Linder P, Friesen N, Bowman JL, Lee JY, Franzke A (2004). Mo-
lecular evidence for bicontinental hybridogenous genomic constitution in Lepidium
sensu stricto (Brassicaceae) species from Australia and New Zealand. Am J Bot 91,
254-261.
28. Lee JY, Mummenhoff K, Bowman JL (2002). Allopolyploidization and evolution of
species with reduced floral structures in Lepidium L.(Brassicaceae). Proc Natl Acad
Sci 99, 16835-16840.
29. Jonsell B, Jarvis CE (2002). Lectotypification of Linnaean names for Flora Nordica
(Brassicaceae – Apiaceae). Nord J Bot 22: 67-86.
30. Marais W (1970). Cruciferae. In L. E. Codd, B. de Winter, D. J. B. Killick, and H. B.
Rycroft (eds.), Flora of Southern Africa vol. 13. Pretoria: Government Printer.
31. Buxbaum JC (1728). Plantarum minus cognitarum centuria1. Typographia Academiae.
32. German DA 2012. Typification of some names in the Cruciferae, mostly from North
Asia. Turczaninowia 15: 9–16.
33. Hedge I, Rechinger KH (1968). Crucifeare. In: Rechinger KH (ed.), Flora Iranica 57.
Graz, Austria: Akademische Druck- u.
34. Mayer J (1786). Abhandlungen der Königlichen Böhmischen Gesellschaft der Wis-
senschaften, ser. 2.
35. Mutlu B (2012). Lepidium L. – In Güner A, Aslan S, Ekim T, Vural M, Babaç MT
(eds.), Türkiye Bitkileri Listesi (Damarlı Bitkiler). Nezahat Gökyiğit Botanik Bahç-
esi ve Flora Araştırmaları Derneği Yayını.
36. Waldstein FC, Kitaibel P (1799). Descriptiones et Icones Plantarum Rariorum Hun-
gariae, Volume 1. Matthiae Andreae Schmidt.
37. Jafri SMH (1973). Flora of West Pakistan No: 55 Brassicaceae. Department of Bot-
any, University of Karachi.
 62 Taxonomic revision of Lepidium L. (Brassicaceae) from Turkey

38. Tournefort JP 1717. Relation d’un voyage du Levant 2.

39. Caferty S, Jarvis CE (2002). Typification of Linnaean plant names in Brassicaceae
(Cruciferae). Taxon 51: 529–537.
40. Ünal M, Özgökçe F, Behçet L, Karabacak O (2007). A new record for Turkey: Lepid-
ium pinnatifidum Ledeb. (Brassicaceae). Turk J Bot 31: 575–576.

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