Turk J Bot
(2015) 39: 76-87
© TÜBİTAK
doi:10.3906/bot-1401-10
Turkish Journal of Botany
http://journals.tubitak.gov.tr/botany/
Research Article
A conspectus of Scorzonera s.l. in Turkey
1
2,
1
2
1
Kamil COŞKUNÇELEBİ , Serdar MAKBUL * , Mutlu GÜLTEPE , Seda OKUR , Murat Erdem GÜZEL
1
Department of Biology, Science Faculty, Karadeniz Technical University, Trabzon, Turkey
2
Department of Biology, Faculty of Sciences and Arts, Recep Tayyip Erdoğan University, Rize, Turkey
Received: 06.01.2014
Accepted: 17.07.2014
Published Online: 02.01.2015
Printed: 30.01.2015
Abstract: A comprehensive taxonomic study based on comparative morphology of Scorzonera, here maintained in its wide sense, is
presented for the territory of Turkey. This study has produced several changes of classification at sectional and species ranks. An updated
list of Scorzonera taxa occurring in Turkey, along with their infrageneric classification, is provided. A new section, S. sect. Anatolia
Makbul & Coskunç., is described. Chromosome numbers, threat categories, and distribution maps are given for 6 endemic taxa (S.
boissieri, S. karabelensis, S. longiana, S. sandrasica, S. ulrichii, and S. zorkunensis) placed in or transferred to the newly described section.
A revised identification key to all Scorzonera species in Turkey is presented.
Key words: Identification key, Podospermum, Pseudopodospermum, Scorzonera, systematics, Turkey
1. Introduction
The genus Scorzonera L. (Asteraceae), with about 160
species belonging to the subtribe Scorzonerinae Dumort.
of the tribe Cichorieae, is widespread in the more arid
regions of Eurasia and northern Africa (Bremer and
Anderberg, 1994; Nazarova, 1997).
After the genus Scorzonera s.str. was revised by
Chamberlain (1975) for the Flora of Turkey, many new
species have been recorded from Turkey. The classification
of the genus has been highly controversial (Nazarova,
1997; Mavrodiev et al., 2004), and because of this different
taxonomic treatments have been used in the floras. Starting
with De Candolle (1805), frequently Scorzonera species with
a conspicuous cylindric carpopodium were placed under a
separate genus, Podospermum DC. (Cassini, 1826; Dumortier,
1827; Endlicher, 1838; Grosheim, 1949; Kuthatheladze,
1978; Pignatti, 1982). Conversely, the genus was maintained
in a wider circumscription but divided into 4 sections
by Boissier (1875), Podospermum, Euscorzonera Lipsch.,
Lasiospora Less., and Epilasia (Bunge) Benth. or divided
into 3 subgenera (Podospermum, Pseudopodospermum
(Lipsch. et Krasch.) Lipsch., and Scorzonera) by Lipschitz
(1935, 1939) and Kamelin and Tagaev (1986). Scorzonera
in a wide circumscription was used in many floras as well,
e.g., Chamberlain (1975) and Chater (1976); Pignatti (1982),
however, was an exception. First phylogenetic analyses of
* Correspondence: smakbul@hotmail.com
76
Scorzonera by Mavrodiev et al. (2004) and Owen et al. (2006)
provided evidence that the previous morphology-based
classifications are at least partly artificial. These 2 analyses
arrived at partly different phylogenetic reconstructions with
respect to the relationships of the major clades. According
to Mavrodiev et al. (2004), Scorzonera, Podospermum, and
Lasiospora should be recognized as distinct genera. The
Turkish species S. rigida and S. seidlitzii, treated under the
sect. Pulvinares by Kamelin and Tagaev (1986), were grouped
together with Lasiospora latifolia Fisch. & C.A.Mey. and L.
hirsuta (Gouan) Cass. with 100% bootstrap value (Mavrodiev
et al., 2004). In addition, the phylogenetic reconstruction
by Owen et al. (2006) revealed that Lasiospora represents a
lineage separate from Scorzonera.
Since the genus Scorzonera in its wide sense was revised
by Chamberlain (1975) to comprise 42 species in Turkey,
many botanists have published new taxa (Coskuncelebi
et al., 2012; Makbul et al., 2012) and new records (Doğan
and Duran, 2010) from Turkey without considering
classification. According to Makbul (2012), the genus
Scorzonera s.l. is represented with 49 species (57 taxa) in
Turkey. Currently, Scorzonera in its wide sense is represented
by 52 species (59 taxa) in Turkey, and among these 31 are
endemics. Recently, some palynological (Blackmore, 1982;
Türkmen et al., 2010), anatomical (Makbul et al., 2011a,
2011b), cytological (Owen et al., 2006; Martin et al., 2012),
and phylogenetic (Mavrodiev et al., 2004) studies have been
COŞKUNÇELEBİ et al. / Turk J Bot
carried out on Turkish Scorzonera. In this comprehensive
study, an updated infrageneric and generic classification,
together with a checklist and new key for Scorzonera taxa
distributed in Turkey, is presented.
2. Materials and methods
This study was based on the examination of herbarium
specimens from the following herbaria: E, K, LE, ANK, EGE,
HUB, GAZI, KNYA, ISTE, ISTF, and ISTO, and the author’s
own specimens collected between 2009 and 2012 from the
country and stored in the herbarium of Karadeniz Technical
University, Department of Biology (KTUB) and the Recep
Tayyip Erdoğan University, Department of Biology (RUB).
Additionally, we have used photographs from the herbaria
B, G, GOET, LINN, and W. All examined specimens (see
Appendix on Journal’s wessite) have been identified by the
first and second authors. In total 739 samples (518 stored
in national herbaria, including our own specimens, and
221 stored in international herbaria) that originated from
Turkey were observed, and several taxonomical names have
been assessed according to all available data.
For karyological analysis, achenes of Scorzonera
boissieri Lipsch., S. hieraciifolia Hayek, S. karabelensis
Parolly & N.Kilian, S. longiana Sümbül, S. sandrasica
Hartvig & Strid, S. ulrichii Parolly & N.Kilian, and S.
zorkunensis Coskunç. & Makbul were germinated in petri
dishes to obtain root tips. Chromosome count was carried
out according to a modified procedure of Coskuncelebi
and Vladimirov (2008).
Threat categories of the taxa belonging to sect. Anatolia
were assessed according to criteria of IUCN (2012) based
on the number of subpopulations and on the size and
quality of the habitats.
3. Results and discussion
3.1. New section
Scorzonera sect. Anatolia Makbul & Coskunç., sect. nov.
Type: Scorzonera zorkunensis Coskunç. & Makbul in
Turkish Journal Bot. 36: 299–310, 2012.
Turkey. C6 Osmaniye: Amanos mountain, Zorkun
plateau, Keldaz hill, 2075 m, 05.07.2010, S.Makbul &
K.Coskuncelebi 242 (holo.: KTUB!, iso.: RUB, ANK!).
Diagnoses: S. sect. Anatolia is similar to sect. Fibrillosae
Nakai but differs from S. sect. Fibrillosae by having brown
dead leaf residues (not fiber-like) at root-collar, densely
hairy leaves (not glabrous), several (not 1–3) basal leaves
crowded near the base, completely yellow ligulate flowers
(not pink and yellow), glabrous and lanate achenes (not
glabrous achenes), and plumose and barbellate pappus
hairs (not plumose and scabrous).
Description: Perennial herbs. Stem scapigerous or
subscapigerous. Root collar covered by brown old leaf
residues, leaves mainly basal, rarely scape with leaflets.
Capitula generally 1, but occasionally 2–5 per stem, ligules
are yellow. Achene glabrous or lanate. Pappus bristles
plumose below, barbellate above.
According to Lipschitz (1935, 1964) subg. Scorzonera is
characterized by cylindrical roots, corniculate phyllaries,
astipitate achenes, and simple or pinnatisect leaves. Other
comparisons of traditionally accepted subgenera within
the Scorzonera are given in Table 1. Kamelin and Tagaev
(1986) reported 22 sections within the subg. Scorzonera.
The newly described section (sect. Anatolia) differs
from the sections described previously within the subg.
Scorzonera by the following characters given in Table 2. In
total, the number of sections represented in Turkey is 13,
including the newly described section (Table 3).
Table 1. Comparison of accepted subgenera within the Scorzonera. Heterogeneous means pericarp consists of both parenchymatic and
sclerenchymatic cells and homogenous means the pericarp consists of only parenchymatic cells.
Characters
Subg. Scorzonera sect. Anatolia
Subg. Podospermum
Subg. Pseudopodospermum
Cylindrical
Cylindrical
Tuberous
Phyllaries
Without corniculate appendage
With corniculate
appendage
Without corniculate appendage
Achene
Non stipitate
Stipitate
Stipitate
Achene surface
Faveolate or smooth and vertical
stripes or rugose
Faveolate and with
horizontal stripes
Faveolate and rugose or vertical
stripes
Lacuna type
Lanata or laciniata
Lanata
Laciniata
Basic chromosome number (X)
6, 7
7
7
Pericarp structure
Heterogeneous
Homogeneous
Heterogeneous
Distribution of vessel in root
Diffuse or radially rays
Radially rays
Diffuse rays
Cross section of the main midrib
V-shaped or elliptic
Triangular
Semicircular
Root
77
COŞKUNÇELEBİ et al. / Turk J Bot
Table 2. Comparison of the sections within the Scorzonera.
Characters
sect. Anatolia
sect. Fibrillosae
sect. Pulvinares
Stem
Mostly subscapigerous
Mostly subscapigerous
Mostly scapigerous
Leaf residues
Not fiber-like
Fiber-like
Not fiber-like
Basal leaves
Several (>3)
1–3
1–3
Leaf indumentum
Densely hairy
Glabrous
Slightly pubescence or subglabrous
Ligules
Yellow
Pink or yellow
Yellow
Achene
Glabrous
Lanate
Glabrous
Pappus
Plumose below, barbellate
above
Plumose below, scabrous above
Plumose below, barbellate above
Habitus
Not caespitose
Not caespitose
Distinctly caespitose
General distribution
Mediterranean element
(endemic to Turkey)
Endemic to Russia and China
Irano-Turanian and Euro-Siberian
element
Six endemic taxa (Scorzonera boissieri, S. karabelensis,
S. longiana, S. sandrasica, S. ulrichii, and S. zorkunensis)
were treated under sect. Anatolia in the present study.
All members of sect. Anatolia share similar micro- and
macromorphological features and are mainly distributed
in South Anatolia (Figure 1). Although an unpublished
phylogenetic analysis by the authors based on sequence
data of nrDNA-ITS and cpDNA supports establishment of
sect. Anatolia under the S. subg. Scorzonera, Mavrodiev et
al. (2004) and Owen et al. (2006) stated that Scorzonera s.l.
is clearly polyphyletic and includes distinct lineages, to be
recognized as separate genera. Mavrodiev et al. (2004) also
reported that these lineages consist of many groups that are
still not sufficiently delimited based on molecular data. On
the other hand, Owen et al. (2006) reported that the genus
Scorzonera s.l. could be divided into 2 distinct groups
based on cytogenetic and molecular data: 1 with 2n = 12
and a second with 2n = 14; however, Nazarova (1997) also
reported that S. subg. Scorzonera includes taxa with 2n =
12 (sect. Nervosae: S. latifolia; sect. Pulvinares: S. seidlitzii,
S. rigida) and 2n = 14 (sect. Nervosae: S. ketzkhovelii)
and does not confirm the division of the genus into the
sections suggested by Lipschitz (1964). Anatolia includes
taxa with 2n = 14 (S. longiana, S. boissieri), 2n (4x) = 28 (S.
zorkunensis), and 2n = 12 (S. karabelensis, S. sandrasica,
S. ulrichii). Our results agree with the view of Nazarova
(1997), but do not concur with the view of Owen et al.
(2006). Mavrodiev et al. (2004) and Owen et al. (2006)
suggested treating Lasiospora as a separate genus including
the Turkish species S. rigida, S. seidlitzii, and S. latifolia;
however, our cytologic and morphologic data of S. rigida
and S. seidlitzii show that these taxa should be placed
under the subg. Scorzonera sect. Pulvinares, as indicated
by Lipschitz (1964) and Nazarova (1997).
78
Members of S. sect. Anatolia generally grow on rockystony slopes in alpine meadows (S. boissieri, S. longiana, S.
zorkunensis) and clearance or boundaries of pine forests
(S. karabelensis, S. ulrichii, S. sandrasica) under the effect
of the Mediterranean climate. Members of this section are
given in an alphabetical order together with the updated
threat category, chromosome number, and other valuable
data as follows.
3.1.1. Scorzonera boissieri Lipsch., Bot. Mater. Gerb. Bot.
Inst. Komarova Akad. Nauk SSSR 22: 293 (1963).
Type: Turkey, B6 Adana: Saimbeyli, Bozoğlan mountain,
above Obruk plateau, 2300 m, Davis, Dodds & Çetik, D.
19751 (holo.: E!).
Chromosome number: 2n = 14 (Makbul &
Coşkunçelebi, 250).
Flowering/fruiting period: May–June/June–July.
Habitat: Clay meadows, slope steppes.
Distribution: Endemic, Southeast Central Anatolia,
Irano-Turanian element.
Reassessed threatened category: VU: B2ab (ii, iii) (VU,
according to Ekim et al., 2000).
This endemic taxon was not previously placed under
any subgenus. The phenetic features supported to place
this taxon under the newly described sect. Anatolia.
Subgeneric classification and chromosome number of S.
boissieri are given here for the first time.
3.1.2. Scorzonera karabelensis Parolly & N.Kilian,
Willdenowia 33: 328 (2003).
Type: Turkey, C2 Muğla: Fethiye–Korkuteli road, below
Karabel pass, NE Kemer, steep, gravely, rocky slopes, open
Pinus nigra Arnold var. caramanica (Loudon) Rehder
forest, N-exp., limestone, 1040 m, 20.05.2003, R. Ulrich
3/7a (holo.: B photo!; iso.: E! ISTE!).
COŞKUNÇELEBİ et al. / Turk J Bot
Table 3. Infrageneric grouping of the Turkish Scorzonera.
Species
S. cana
S. armeniaca
S. laciniata
S. hieraciifolia
S. aksekiensis
S. elata
S. inaequiscapa
S. suberosa
S. phaeophappa
S. mollis
S. semicana
S. boissieri
S. karabelensis
S. longiana
S. sandrasica
S. ulrichii
S. zorkunensis
S. acuminata
S. davisii
S. pacis
S. incisa
S. lacera
S. violacea
S. acantholimon
S. argyrea
S. cinerea
S. ketzkhovelii
S. mirabilis
S. veratrifolia
S. bella
S. latifolia
S. dzhawakhetica
S. tomentosa
S. papposa
S. parviflora
S. tuzgoluensis
S. amasiana
S. lasiocarpa
S. sericea
S. seidlitzii
S. rigida
S. pygmaea
S. yildirimlii
S. ahmet-duranii
S. coriacea
S. aucheriana
S. sublanata
S. pseudolanata
S. renzii
S. pisidica
S. kotschyi
S. eriophora
According to cited literature
Present study
Subg. Podospermum (Lipschitz, 1964)
Subg. Podospermum (Lipschitz, 1964)
Subg. Podospermum (Lipschitz, 1964)
Subg. S. sect. Pseudopodospermum (Kamelin & Tagaev, 1986)
Subg. Podospermum
Subg. Podospermum
Subg. Podospermum
Subg. Podospermum
Subg. S. sect. Foliosae (Kamelin & Tagaev, 1986)
Subg. S. sect. Papposae (Kamelin & Tagaev, 1986)
Subg. Pseudopodospermum (Lipschitz, 1964)
Subg. S. sect. Pseudopodospermum (Kamelin & Tagaev, 1986)
Subg. Pseudopodospermum (Lipschitz, 1964)
Subg. S. sect. Pseudopodospermum (Kamelin & Tagaev, 1986)
Subg. Pseudopodospermum
Subg. Pseudopodospermum
Subg. Pseudopodospermum
Subg. Pseudopodospermum
Subg. Pseudopodospermum
Subg. Pseudopodospermum
Subg. Pseudopodospermum
Subg. S. sect. Nervosae (Parolly & Kilian, 2003)
Subg. S. sect. Nervosae (Parolly & Kilian, 2003)
Subg. S. sect. Nervosae (Parolly & Kilian, 2003)
Subg. S. sect. Nervosae (Parolly & Kilian, 2003)
Subg. S. sect. Nervosae (Parolly & Kilian, 2003)
Subg. S. sect. Nervosae (Coskuncelebi et al., 2012)
Subg. S. sect. Foliosae (Kamelin & Tagaev, 1986)
Subg. S. sect. Foliosae (Kamelin & Tagaev, 1986)
Subg. S. sect. Anatolia
Subg. S. sect. Anatolia
Subg. S. sect. Anatolia
Subg. S. sect. Anatolia
Subg. S. sect. Anatolia
Subg. S. sect. Anatolia
Subg. S. sect. Foliosae
Subg. S. sect. Foliosae
Subg. S. sect. Foliosae
Subg. S. sect. Incisae (Kamelin & Tagaev, 1986)
Subg. S. sect. Incisae (Kamelin & Tagaev, 1986)
Subg. S. sect. Incisae (Kamelin & Tagaev, 1986)
Subg. S. sect. Incisae
Subg. S. sect. Incisae
Subg. S. sect. Incisae
Subg. S. sect. Infrarosulares (Kamelin & Tagaev, 1986)
Subg. S. sect. Infrarosulares
Subg. S. sect. Nervosae (Kamelin & Tagaev, 1986)
Subg. S. sect. Nervosae (Kamelin & Tagaev, 1986)
Subg. S. sect. Nervosae (Kamelin & Tagaev, 1986)
Subg. S. sect. Nervosae (Kamelin & Tagaev, 1986)
Subg. S. sect. Nervosae (Kamelin & Tagaev, 1986)
Subg. S. sect. Nervosae (Kamelin & Tagaev, 1986)
Subg. S. sect. Nervosae
Subg. S. sect. Nervosae
Subg. S. sect. Nervosae
Subg. S. sect. Nervosae
Subg. S. sect. Nervosae
Subg. S. sect. Nervosae
Subg. S. sect. Nervosae
Subg. S. sect. Nervosae
Subg. S. sect. Nervosae
Subg. S. sect. Nervosae (Lipschitz, 1964; Kamelin & Tagaev, 1986)
Subg. S. sect. Nervosae (Lipschitz, 1964; Kamelin & Tagaev, 1986)
Subg. S. sect. Gelasia (Kamelin & Tagaev, 1986)
Subg. S. sect. Papposae (Lipschitz, 1964; Kamelin & Tagaev, 1986)
Subg. S. sect. Parviflorae (Lipschitz, 1964; Kamelin & Tagaev, 1986)
Subg. S. sect. Pulvinares (Kamelin & Tagaev, 1986)
Subg. S. sect. Pulvinares (Kamelin & Tagaev, 1986)
Subg. S. sect. Pulvinares (Kamelin & Tagaev, 1986)
Subg. S. sect. Pulvinares (Lipschitz, 1964)
Subg. S. sect. Pulvinares (Lipschitz, 1964; Kamelin & Tagaev, 1986)
Subg. S. sect. Pulvinares (Kamelin & Tagaev, 1986)
Subg. S. sect. Subaphyllae (Kamelin & Tagaev, 1986)
Subg. S. sect. Tuberosae (Kamelin & Tagaev, 1986)
Subg. S. sect. Tuberosae (Lipschitz, 1964)
Subg. S. sect. Turkestanicae (Kamelin & Tagaev, 1986)
Subg. S. sect. Nervosae (Parolly and Kilian, 2003)
Subg. S. sect. Vierhapperia (Kamelin & Tagaev, 1986)
Subg. S. sect. Vierhapperia (Kamelin & Tagaev, 1986)
Subg. S. sect. Papposae
Subg. S. sect. Parviflorae
Subg. S. sect. Parviflorae
Subg. S. sect. Pulvinares
Subg. S. sect. Pulvinares
Subg. S. sect. Pulvinares
Subg. S. sect. Pulvinares
Subg. S. sect. Pulvinares
Subg. S. sect. Pulvinares
Subg. S. sect. Pulvinares
Subg. S. sect. Scorzonera
Subg. S. sect. Scorzonera
Subg. S. sect. Subaphyllae
Subg. S. sect. Tuberosae
Subg. S. sect. Tuberosae
Subg. S. sect. Turkestanicae
Subg. S. sect. Vierhapperia
Subg. S. sect. Vierhapperia
Subg. S. sect. Vierhapperia
79
COŞKUNÇELEBİ et al. / Turk J Bot
Figure 1. Distribution map of sect. Anatolia in Turkey.
Chromosome number: 2n = 12 (first report in this
study) (Makbul 134).
Flowering/fruiting period: May–June/June–August.
Habitat: Rocky-stony slopes, clearance of Pinus nigra
forest.
Distribution: Endemic, South Anatolia, Mediterranean
element.
Reassessed threatened category: VU: B2ab (iii, iv) (VU,
according to Parolly and Kilian, 2003). This local endemic
was recollected from Karabel Geçidi (type locality), and
according to Parolly and Killian (2003) its distribution
reaches the upper slopes of Akdümen Tepe (1742 m) in the
greater Boncuk mountain range. The habitat quality and
the number of mature individuals lead us to treat this taxa
under VU according to IUCN (2012).
This stenoendemic taxon was previously treated under
subg. Scorzonera sect. Nervosae Lipsch. by Parolly and Kilian
(2003). According to Lipschitz (1935, 1964), sect. Nervosae
is characterized by large perennials with densely leafy
stems and tomentose indumentums, but S. karabelensis is a
scapigerous to subscapigerous perennial herb with sericeous
appressed indumentums. Hence, it is more proper to place
S. karabelensis under the sect. Anatolia based on the gross
morphological characters given above.
3.1.3. Scorzonera longiana Sümbül, Edinburgh J. Bot. 48:
35 (1991).
Type: Turkey, C4 Antalya: Gazipaşa, from Çobanlar village
plateau to Oyuklu plateau, 1900–2000 m, 11.07.1983,
Sümbül 2239 (holo.: HUB!, iso.: E! K photo!).
Chromosome number: 2n = 14 (first report in this
study) (Makbul & Coşkunçelebi, 255).
Flowering/fruiting period: July/July–August.
Habitat: Alpine, stony meadows.
Distribution: Endemic, South Anatolia, Mediterranean
element.
Reassessed threatened category: EN: B2ab (ii, iii) (EN,
according to Ekim et al., 2000).
Scorzonera longiana was not previously placed into
any infrageneric group. The phenetic features support its
placement in S. sect. Anatolia of S. subg. Scorzonera.
80
3.1.4. Scorzonera sandrasica Hartvig & Strid, Bot. Jahrb.
Syst. 108: 311 (1987).
Type: Turkey, C2 Muğla: Sandras mountain, W of the
summit, slightly sloping stony flat (snowbed) near a small
creek close to timberline, serpentine, 1970 m, 07.07.1984,
N 37 04-E 028 50, P. Hartvig, Ö. Seçmen & A. Strid 23342
(holo.: C; iso.: E!).
Chromosome number: 2n = 12 (first report in this
study) (Makbul & Coşkunçelebi, 232).
Flowering/fruiting period: June–July/July–August.
Habitat: Above forest boundaries, alpine hills, rocky
grasslands.
Distribution:
Endemic,
Southwest
Anatolia,
Mediterranean element.
Reassessed threatened category: EN: B2ab (i, ii) (EN,
according to Ekim et al., 2000).
This endemic taxon was not previously placed into any
infrageneric group. It is only known from the type locality,
and its morphological features suggest its placement in the
newly described S. sect. Anatolia.
3.1.5. Scorzonera ulrichii Parolly & N. Kilian, Willdenowia
32: 198 (2002).
= Scorzonera gokcheoglui Ünal & Göktürk, Bot. J. Linn.
Soc. 142: 465 (2003).
Type: Turkey, C4 Antalya: Distr. Alanya, Mahmutlar–
Hadim road, c. 35 km NE of Mahmutlar and 15 km S of
Çayarası, 1 km N of Elmalısu, gravely, rocky slopes with
clearance of Pinus nigra var. caramanica forest, W-exp.,
limestone, 1250 m, 11.06.2002, Robert Ulrich (Tubingen)
2/12 (holo.: B photo!; iso.: E! ISTE!).
Chromosome number: 2n = 12 (first report in this
study) (Makbul & Coşkunçelebi, 237).
Flowering/fruiting period: June–July/July.
Habitat: Slopes, clearance of Pinus, calcareous and
stony areas.
Distribution: Endemic, South Anatolia, Mediterranean
element.
Reassessed threatened category: VU: B2ab (ii, iv) (EN,
according to Parolly and Kilian, 2002).
COŞKUNÇELEBİ et al. / Turk J Bot
This endemic species was previously placed under
the S. subg. Scorzonera sect. Nervosae. The section is
recognized for its large perennials with densely leafy
stems and tomentose indumentum as stated by Parolly
and Kilian (2003). However, S. ulrichii is a scapigerous
to subscapigerous caespitose perennial herb with lanate
indumentum. Therefore, it is more appropriate to place it
under the sect. Anatolia.
3.1.6. Scorzonera zorkunensis Coskunç. & Makbul, Turk.
J. Bot. 36 (4): 302 (2012).
Type: Turkey, C6 Osmaniye: Amanos mountains, 5 km
after Zorkun plateau, Keldaz hill, Halepgören area, 2075 m,
05.07.2010, Makbul & Coskuncelebi, 242 (holo.: KTUB!;
iso.: RUB! ANK!).
Chromosome number: 2n = 4x = 28 (first report in this
study) (Makbul & Coşkunçelebi, 242).
Flowering/fruiting period: July/July–August.
Habitat: Serpentine steppe.
Distribution:
Endemic,
South-East
Anatolia,
Mediterranean element.
Reassessed threatened category: EN: B2a (ii, iii) (EN,
according to Coskuncelebi et al., 2012).
Scorzonera zorkunensis was previously treated
under the subg. Scorzonera sect. Nervosae as stated by
Coskuncelebi et al. (2012). Scorzonera zorkunensis is a
scapigerous and mostly caespitose perennial herb with
lanate indumentum. Thus, we decided that it is more
suitable to treat it under the sect. Anatolia.
3.2. New synonyms
3.2.1. Scorzonera davisii Lipsch., Bot. Mater. Gerb. Bot.
Inst. Komarova Akad. Nauk SSSR, 22: 291 (1963).
Type: Turkey, C9 Hakkari: Zap gorge, 48 km S of Başkale,
03.08.1954, Davis 23848 & Polumin (holo.: E!; iso.: K
photo!).
= Scorzonera tenax Rech. f., Fl. Iranica 122:59 (1977),
syn. nov.
Type: Distr. Mosul (Kurdistan), 10.07.1957, Rechinger
11572 (holo.: B photo!); ad confines Turciae prov. Hakkari,
inter Dohuk et Amadiya; in quercetis saxosis supra Suwara
Tuka.
After a detailed examination of the specimens
stored in E (E00385297! for S. davisii Lipsch. and
E00476936!-E00224242!-E00224241! for S. tenax Rech.
f.) and comparison of the specimens described in the
original papers (Lipschitz, 1964; Rechinger, 1977), we
determined that the 2 taxa are very similar in terms of
some morphological features such as entire leaves and
glabrous and verrucose achenes and that they fall within
the variation of S. davisii. Based on all available evidence,
we conclude that S. tenax is conspecific of S. davisii.
3.2.2. Scorzonera rigida DC., Prodr. 7: 123 (1838).
Type: Turkey, B7 Nevşehir: the high mountains of
Cappadocia and the Euphrates, Aucher 3309 (holo.: G DC
photo!; iso.: K!).
= Scorzonera adilii A.Duran, Pakistan J. Bot. 34: 385
(2002), syn. nov.
Type: C5 Konya: Ereğli, Aydos mountain, Kayasaray
village, Düğünlük stream, 1770 m, crevices of marble
rocks, 07.07.2001, N 3722.04-E 3416.33, A. Duran 5798
& Sağıroğlu (holo.: ADO; iso.: GAZI! ANK! HUB!), ibid.,
18.07.2002, A.Duran 5960 (ADO).
= Scorzonera aytatchii A.Duran & Sağiroğlu, Nordic J.
Bot. 22: 333 (2003).
Type: C5 Konya: Ereğli, Aydos mountain, Kayasaray
village, Düğünlük stream, 1750 m, crevices of marble
rocks, 07.07.2001, N 3722-E 3416, A. Duran 5797 &
Sağıroğlu (holo.: ADO, iso.: GAZI! ANK! HUB!); ibid.
Sayıntaş area, 15.07.1977, S. Erik 2602 (para.: HUB).
S. adilii is described from Aydos mountain, Konya,
Turkey (Duran, 2002). The features of S. adilii reported
by Duran (2002) are not sufficiently clear to separate this
taxon from S. rigida. Entirely barbellate pappus hairs are
also seen in some specimens of S. adilii, as in S. rigida.
These 2 taxa are morphologically very similar. Both taxa
are subscapigerous perennial herbs with caespitose habitus
and have short-pilose achene indumentums. According to
Chamberlain (1975) and Parolly and Kilian (2003), S. rigida
is a highly variable taxon distributed mainly throughout
SE and E Anatolia, but it also appeared in Aydos mountain
(Konya). S. adilii was discovered from the adjacent area
of the type locality of S. aytatchii A.Duran & Sağıroğlu
(2002), which Parolly and Kilian (2003) named S. rigida.
The range of variation observed in both quantitative and
qualitative characters of S. adilii fall fully within S. rigida.
Thus, S. adilii was considered a synonym of S. rigida.
3.2.3. Scorzonera hieraciifolia Hayek, Ann. K. K.
Naturhist. Hofmus. 20: 426 (1907).
Type: Turkey, B5 Kayseri: Erciyes mountain, in der
Salzsteppe bei Soisaly, c. 1100 m, v 1902, Zederbauer
(holo.: W photo!).
= Scorzonera kurtii Yıldırımlı, Ot Sistematik Botanik
Dergisi 18 (2): 12 (2011), syn. nov.
Type: Turkey, B4 Konya: Kulu, Tuzyaka (Sütkanlı),
Bozan village, Kaldırım road, Tuzgölü coast, salty places,
945 m, 04.05.2007, Ş.Yıldırımlı 33661 (holo.: Yıldırımlı
otluk; iso.: HUB!).
Scorzonera hieraciifolia is highly variable taxon
distributed mainly in salty habitats around Tuzgölü
(Konya). Scorzonera hieraciifolia shows a range of
variation observed in the leaves and stem. Scorzonera
kurtii is separated from S. hieraciifolia (Yıldırımlı, 2012)
mainly by leaf and stem features that are in the range of
variation observed in the subpopulation of S. hieraciifolia
81
COŞKUNÇELEBİ et al. / Turk J Bot
distributed around Tuzgölü. In particular, pinnatisect
leaves, achene characteristics, and salty habitus of the S.
kurtii are very similar to some specimens of S. hieraciifolia
stored in RUB (Makbul 127!, 138!). Additionally, the range
of variation observed in both quantitative and qualitative
characters of S. kurtii fall fully within S. hieraciifolia. Thus
S. kurtii was considered a synonym of S. hieraciifolia.
3.2.4. Scorzonera pygmaea Sibth. & Sm., Fl. Graec. Prodr.
2: 122 (1813).
Type: Turkey, A2 (A) Bursa, Olympo Bithyno (Uludağ),
cacumine, Sibthorp.
Lectotype (designated here): Turkey, A2 Bursa,
Uludağ, 2380–2580 m, 13 ix 1949, D. 14846 (E!).
= S. pygmaea Sibth. & Sm. subsp. nutans (Czeczott)
Chamberlain, Notes R.B.G.: Edinb. 33:433 (1975) ≡ S.
pygmaea Sibth. & Sm. var. nutans (Czeczott) O.Koyuncu
& Yaylacı, Pakistan Journal of Botany, 45, 5 (2013) ≡
Scorzonera nutans Czeczott in Repert. Spec. Nov. Regni
Veg. Beih. 107: 202. 1938, syn. nov.
Further examinations of many recent samples from
type localities of S. pygmaea subsp. pygmaea and S. pygmaea
subsp. nutans and type specimens stored in E herbarium
did not support the separation of these 2 taxa as indicated
by Chamberlain (1975). Many samples including the types
and recent collections share similar ranges in stem and leaf
size. In addition, detailed gross morphological examination
of specimens belonging to S. pygmaea subsp. pygmaea and
S. pygmaea subsp. nutans revealed that these 2 taxa are
very similar in terms of character combination of general
habitus, phyllary, and achene shape and size. Additionally,
our phylogenetic analysis based on unpublished
molecular data (submitted to GenBank) belongs to several
populations of both S. pygmaea subsp. pygmaea (GenBank
accession no.: KF925533) and S. pygmaea subsp. nutans
(GenBank accession no.: KF925532) and supports the
integration of subspecies under S. pygmaea. Parolly and
Kilian (2003) reported that it is not possible to separate
these 2 taxa based on the traits referred to by Chamberlain
(1975). Recently, Koyuncu et al. (2013) suggested that
these 2 subspecies should be described as varieties because
of the similarity in traits and the overlapping distribution
range. However, Koyuncu et al. (2013) did not give enough
evidence for separating these taxa. Additionally, Koyuncu
et al. (2013) reported that there are no distinct differences
among the examined specimens based on anatomical,
palynological, and ecological features. Hence, we decided
that S. pygmaea var. pygmaea, and S. pygmaea var. nutans
should be combined under the name of S. pygmaea.
3.3. New suggestions for the subgeneric level
After detailed examination of several specimens of
Scorzonera hieraciifolia, which is treated under subg.
82
Podospermum (DC.) Lipsch. sect. Pseudopodospermum
Lipsch. (Kamelin and Tagaev, 1986), we decided that
S. hieraciifolia should be placed in subg. Podospermum
(DC.) Lipsch. because of the stipitate achene, corniculate
phyllaries, and cylindrical roots. Although Mavrodiev et
al. (2004) and Owen et al. (2006) proposed that the genus
Podospermum be recognized as separate genera, they did
not supply sufficient molecular data. The present authors
follow Lipschitz (1935, 1939) and Kamelin and Tagaev
(1986). Additionally, some unpublished data obtained
from achene ultrastructure, basic chromosome number,
and pollen lacuna types, according to Blackmore (1982),
suggest S. hieraciifolia be placed under subg. Podospermum.
Scorzonera hieraciifolia has lanata type pollen which is the
common pollen type of subg. Podospermum (Blackmore,
1982). All examined members of Podospermum were
characterized by a pericarp, which consisted of only
sclerenchymatic cells (homogenous pericarp). It was
determined that S. hieraciifolia has a homogenous pericarp
structure as in subg. Podospermum. Additionally, the
chromosome number of S. hieraciifolia is 2n = 14 (Makbul
& Coşkunçelebi, 127). According to Nazarova (1997), the
basic chromosome number is x = 7 in all Podospermum
members. Therefore, S. hieraciifolia was transferred to
subg. Podospermum.
Scorzonera elata Boiss. and S. inaequiscapa Boiss.
treated under subg. Scorzonera sect. Foliosae (Boiss.)
Lipsch. and sect. Papposae Lipsch. et Krasch. (Kamelin
and Tagaev, 1986), respectively, were moved to subg.
Pseudopodospermum based on examination of the gross
morphology of several samples. According to Lipschitz
(1935), Pseudopodospermum is characterized by stipitate
achene, non-corniculate phyllaries, and simple leaves.
These characteristics were observed in all specimens of
S. elata and S. inaequiscapa. Corroborating these results,
our further micro-/macromorphological observation
showed that both taxa should be moved to subg.
Pseudopodospermum.
Additionally, S. tomentosa L., treated under S. subg.
Scorzonera sect. Gelasia (Caas.) Less. by Kamelin and
Tagaev (1986), should be moved to subg. Scorzonera sect.
Nervosae Lipsch. Section Gelasia consists of height and
pubescent and caulescent plants with smooth achenes,
but S. tomentosa has densely pubescent caulescent plants
with distinctly ridged achene. Consequently, S. tomentosa
shared specific features with sect. Nervosae, and because of
this it should be moved from sect. Gelasia to sect. Nervosae.
Makbul et al. (2011a, 2011b) reported that stem and
leaf anatomical and palynological properties (Türkmen
et al., 2010) also supported the conventional taxonomic
COŞKUNÇELEBİ et al. / Turk J Bot
treatment suggested by Kamelin and Tagaev (1986).
According to detailed examination of our own and type
specimens (E, RUB, and KTUB), as well as unpublished
cytological, anatomical, palynological data, the taxonomic
rank of S. phaeopappa and S. semicana DC., traditionally
treated under subg. S. sect. Pseudopodospermum
(Kamelin and Tagaev, 1986), was evaluated as S. subg.
Pseudopodospermum (Lipsch. et Krasch.) Lipsch.
Duran et al. (2013) and Güzel et al. (2013) recently
published 2 Scorzonera taxa, S. aksekiensis A.Duran
& M.Öztürk and S. pacis Guzel, Kayikci & S.Yildiz,
respectively, from Turkey. However, they did not suggest
any subgeneric or sectional groups for these 2 taxa.
According to the description appearing in the original
paper, S. aksekiensis should be treated under subgenus
Pseudopodospermum based on stipitate achene, tuberous
root, and entire leaves, and this has been carried out
for the first time in the present study. Similarly, S. pacis
should be treated under subgenus Scorzonera sect. Foliosae
due to the high foliated stem and distinct toothed and
winged achenes. Other infrageneric and sectional changes
suggested here are given in Table 3.
3.4. Excluded and included taxa from the list of
Scorzonera recorded from Turkey
Scorzonera hispanica was recorded for Turkey based on
samples from vegetable sellers at Pera (İstanbul); however,
information about the origin of these samples is missing
from both national and international herbariums. For this
reason we thought that the samples sold by local sellers
belong to S. mollis subsp. mollis. During this review,
intense field work was conducted around Tekirdağ and
Mürefte districts where S. hispanica is possibly distributed
and/or recorded from. We were not able collect or observe
S. hispanica. Nevertheless, populations of S. mollis subsp.
mollis are abundant in the cited area. The individuals of
S. mollis subsp. mollis appear similar, at first glance, to S.
hispanica. Because we thought that S. hispanica may have
been confused with S. mollis or recorded by mistake, this
taxon was excluded from the list of Scorzonera distributed
in Turkey.
Scorzonera cretica Willd. naturally distributed on
Crete (Greek island) was excluded from the list of Turkish
Scorzonera. Intensive field work was carried out to find
this species in Turkey, but all efforts resulted in nothing.
Scorzonera woronowii Krasch recorded from Turkey
by Erik (1990) was based on samples stored in HUB
(31446!, 31447!). However, detail examination of these
samples showed that they belong to S. ketzkhovelii Grossh,
recorded from Turkey by Hamzaoğlu et al. (2010). Thus S.
woronowii was also excluded from the list presented in the
following key.
Scorzonera bella Lipsch. was first described by Lipschitz
(1963) based on the specimens of P.H. Davis & O.V.
Polunin, 23925 (holo LE!, iso E!) collected from Turkey,
Hakkari, Cilo mountain, Diz stream in 1954. However, later
it was verified by Chamberlain (1975) under S. veratrifolia
Fenzl in the Flora of Turkey. A detailed examination of
both types of S. veratrifolia stored in K (K000797281!) and
types of S. bella stored in LE! (holo) and E! (iso) shows
that S. bella is a separate/distinct species. Thus, S. bella was
added to the list of Scorzonera taxa by the authors in the
present study.
3.5. A new identification key to Scorzonera taxa
distributed in Turkey
This key is mainly based on field observations and the
examination of samples collected by the authors and
specimens stored in the national and international herbaria
listed in materials and methods. Chamberlain (1975) gave
a key in his account, without infrageneric grouping, for
39 species, and Parolly and Kilian (2003) gave a key for
the only subscapigerous taxa (23 taxa). In the present
study, we provided a revised and updated identification
key including 52 (59 taxa) species, 31 of them endemic to
Turkey. The distinctive characters used in the identification
key are given in Figures 2 and 3.
1-
−
2-
−
3−
4-
−
5-
−
6−
7-
Achene at base with hollow swollen stipe (stipitate
achene), involucral bracts at apex with or without
corniculate appendages..................................................2
Achene characters not as above (1. Subg. Scorzonera)...12
Rootstock cylindrical, all leaves (stem and basal
leaves) or at least some leaves pinnatisect or deeply
lobed, involucral bracts at apex with corniculate
appendages (2. Subg. Podospermum) ..........................3
Rootstock tuberous, all leaves entire, involucral bracts
at apex without corniculate appendages (3. Subg.
Pseudopodospermum) ...................................................6
Achene lanate; at least mature achene lanate ..............4
Achene glabrous ............................................................5
Some of basal leaves pinnatisect, entire leaves ovate or
elliptic, inner phyllaries 12–15 mm in length, achene
4–7 mm in length.............................18. S. hieraciifolia
Basal leaves pinnatisect, segments linear-lanceolate,
inner phyllaries 15–25 mm in length, achene 9–13
mm in length.........................................7. S. armeniaca
(3) Plants annual or biennial, not crowned by
leaf remains at base, ligules as long as inner
phyllaries..................................................25. S. laciniata
Plants perennial, crowned by leaf remains,
ligules at least 1.5 times as long as inner
phyllaries........................................................11. S. cana
(2) Flowers purple or violet ..........................................7
Flowers yellow ................................................................8
Plants subscapigerous, achene (16–)18–21 mm,
pappus hairs barbellate .........................44. S. suberosa
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COŞKUNÇELEBİ et al. / Turk J Bot
Figure 2. Types of roots and leaves observed in Scorzonera. a- tuberous root, b- cylindrical root, cpinnatifid leaf, d- lanceolate leaf, e- ovate leaf, f- linear leaf, g- plane leaf margin, h- undulate leaf margin.
Figure 3. Valuable characters used in the key of the Turkish Scorzonera. Phyllary (a- aristate, bacuminate, c- acute, d- corniculate appendage); achene (e- lanate, f- glabrous and nonstipitate, gstipitate, h- smooth, i- lamellate, j- verrucose); pappus (k- plumose, l- barbellate, m- scabrid).
84
COŞKUNÇELEBİ et al. / Turk J Bot
−
8−
9-
−
10−
11-
−
1213-
-
14-
-
15−
16−
17−
18−
Plants subscapigerous or caulescent, achene
14–16(–18) mm, end of the pappus hairs
scabrid................................................34. S. phaeopappa
(6) Plants clearly caulescent, branched, leaves slightly
undulate or not, achene 8–12 mm in length ................9
Plants subscapigerous or subcaulescent, rarely
branched or not, leaves densely undulate, achene
15–30 mm in length ....................................................10
Plants 30–95 cm length, glabrous, branches erect,
leaves linear, 1–2(–3)-mm wide, outer and inner
phyllaries acute, achene lamellate ...............16. S. elata
Plants 15–30(–35) cm in length, glabrous or crispedpubescent below, branches ascending to erect, leaves
lanceolate to oblong-lanceolate, 8–12(–15)-mm wide,
outer phyllaries acute to acuminate, inner phyllaries
acuminate, achene muricate ............... 4. S. aksekiensis
(8) Tuber slightly elongated ellipsoid, outer phyllaries
acuminate...............................................42. S. semicana
Tuber spherical, outer phyllaries acute ........................11
Leaves 2–5-mm wide, flowering capitula 22–45
mm in length, outer phyllaries 13–19 mm, achene
lamellate, 15–21 mm..................................30. S. mollis
Leaves 3–14-mm wide, flowering capitula 18–20(25)
mm in length, outer phyllaries 7–12 mm, achene
7–15(17) mm, tuberculate..............19. S. inaequiscapa
(1) At least some basal leaves deeply lobed to
pinnatifid or pinnatisect ............................................ 13
All leaves entire ............................................................15
Capitula (35–)40–50 mm, flowers yellow with purple
tube or with purple stripes lower surface of ligules,
achenes (12–)15–23 mm, pappus hairs barbellate
above, plumose below..................................20. S. incisa
Capitula 30–40 mm, flowers purple to violet, achenes
8–15 mm, some of the pappus hairs barbellate, others
barbellate above, plumose below ...............................14
Plants 15–40 cm, leaves herbaceous, outer phyllaries
14–16 mm, achenes 8–10 mm, surface lamellate ......
.......................................................................24. S. lacera
Plants 6–15 cm, leaves coriaceous, outer phyllaries
7–8 mm, achenes 10–15 mm, surface verrucose ......
...................................................................50. S. violacea
(12) Leaves ovate, elliptic or elliptic-lanceolate, leaves
5 times as long as broad ..............................................16
Leaves linear or linear-lanceolate, leaves more than 5
times as long as broad .................................................25
Achene glabrous .........................................................17
Achene lanate ...............................................................22
Leaves villous or pannose ...........................................18
Leaves tomentose, canescent-pannose or crispedlanate .............................................................................20
Plants <15 cm, ascending, densely pannose, leaves
ovate or obovate, rarely elliptic .................6. S. argyrea
Plants 15–50 cm, erect, villous, leaves lanceolate or
elliptic-lanceolate ........................................................19
19- Achene 10–12 mm, inner phyllaries (14–)15–20 mm
in length......................................................12. S. cinerea
− Achene <10 mm, inner phyllaries 12–15 mm in
length ......................................................29. S. mirabilis
20- (17) Plant subscapigerous, achene 14–17 mm,
verrucose..................................................10. S. boissieri
− Plant caulescent, achene 9–14 mm, smooth ..............21
21- Plants tomentose, outer phyllaries aristate, pappus
red-brown, barbellate ........................46. S. tomentosa
Plants canescent-pannose, outer phyllaries obtuse or
acute, pappus yellowish-cream, plumose-barbellate ..
...........................................................22. S. ketzkhowelii
22- (16) Plants greenish, glabrous or sparsely
villous.........................................................27. S. latifolia
Plants gray or grayish, canascent-lanate, pannose or
lanate..............................................................................23
23- Stem erect, c. 1–10 capitulate, fruiting capitula 30–40
mm in length ...............................15. S. dzhawakhetica
Stem ascending to erect, capitula number more than
10, fruiting capitula <30 mm in length .................... 24
24- Stem distinctly branched especially above,
inflorescence racemose, capitula 18–25 mm in length
..............................................................49. S. veratrifolia
−
Stem unbranched, inflorescence panicula, capitula
<18 mm in length ...........................................9. S. bella
25- (15) Plants suffruticose at base.........1. S. acantholimon
− Plants herbaceous.........................................................26
26- Plants scapigerous, subscapigerous, or subcaulescent
........................................................................................27
Plants caulescent, clearly branched............................43
27- Caespitose plants, leaves linear with a plane margin
........................................................................................28
Plant not caespitose, leaves linear-lanceolate or
lanceolate, sometimes with undulate margin.............33
28- Plants gray, densely sericeous....................42. S. sericea
Plants green, crisped pubescent or lanate....................29
29- Plants (10–)15–30 cm, erect ....................39. S. rigida
Plants 5–10(–15) cm, at least some stems ascending
........................................................................................30
30- Achene lanate......................................26. S. lasiocarpa
Achene glabrous............................................................31
31- All leaves (stem and basal leaves) toward the base
densely lanate, outer phyllaries (6.5–)7–9 mm, lanate
...................................................................41. S. seidlitzii
All leaves (stem and basal leaves) glabrous, sparsely
lanate or crisped, outer phyllaries 3–7 mm, glabrous
or crisped pubescent ...................................................32
32- Outer phyllaries glabrous, aristate ......51. S. yildirimlii
Outer phyllaries crisped pubescent, acute ...................
..................................................................37. S. pygmaea
33- (27) Rootstock creeping. Stems hollow ........................
................................................................33. S. parviflora
Characters not as above ..............................................34
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34- Rootstock tuberous .....................................................35
Rootstock cylindrical .................................................36
35- Scape 12–25 mm, leaves 2–5(–6)-mm wide, plane or
undulate ................................................45. S. sublanata
Scape 5–12 mm, leaves 5–15-mm wide, undulate ......
..........................................................36. S. pseudolanata
36- (34) Leaves coriaceous, achene 16–22 mm .................
...................................................................13. S. coriacea
Leaves not coriaceous, achene shorter than 16 mm ...37
37- Achene lanate ...............................................................38
Achene glabrous............................................................41
38- Outer phyllaries grayish-white, densely lanatepannose, pappus whitish-cream .....21. S. karabelensis
Outer phyllaries greenish, lanate, crisped pubescent or
sericeous, pappus purple..............................................39
39- Plants 5–15 cm, leaves lanate, hairs 3–7 mm in length,
inner phyllaries 14–18 mm, achene 6–9 mm in length
.....................................................................48. S. ulrichii
Plants 2–5(–6) cm, leaves crisped-lanate or sericeous,
hairs shorter than 2 mm, inner phyllaries 10–14 mm,
achene 3–6(–7) mm.....................................................40
40- Leaves 3–5(–6) × 0.2–0.3 cm, densely undulate,
sparsely crisped-lanate, outer phyllaries acuminate ...
...............................................................40. S. sandrasica
Leaves 1–3 × 0.3–0.9 cm, slightly undulate, sericeous,
outer phyllaries acute.............................28. S. longiana
41- (37) Capitula 25–35 mm in length, outer phyllaries
winged, 11–14 mm in length ........3. S. ahmet-duranii
Capitula 15–25 mm in length, outer phyllaries not
winged, 4–11(–12) mm in length...............................42
42- Leaves lanate-pannose, veins prominent, inner
phyllaries 20–22 mm, achene 8–13 mm in length ......
....................................................................35. S. pisidica
Leaves lanate, veins not prominent, inner phyllaries
9–20 mm, achene 15–21 mm in length .....................
............................................................51. S. zorkunensis
43- (26) Flowers purple ...............................32. S. papposa
Flowers yellow................................................................44
44- Achene lanate ..............................................................45
Achene glabrous...........................................................47
45- Plants lanate-pannose, leaves 10–35-mm wide,
capitula 25–40 mm in length, outer phyllaries 5–10
mm, achene 8–10 mm in length...........17. S. eriophora
Plants sparsely lanate, leaves 2–10-mm wide, capitula
8–25 mm in length, outer phyllaries 3–5(–6) mm,
achene 4–6 mm in length ...........................................45
46- Plants ascending, leaves 2–4-mm wide, outer
phyllaries obtuse-acute...........................23. S. kotschyi
Plants erect, leaves 5–8-mm wide, outer phyllaries
acuminate...............................................8. S. aucherana
47- (44) Leaves subcoriaceous, 10–18-mm wide ...............
.................................................................2. S. acuminata
Leaves not as above, 1–10-mm wide..........................48
48- Achene 17–20 mm in length....................14. S. davisii
Achene shorter than 15 mm.......................................49
49- Caespitose plants branched at base, villous, mature
capitula 15–20(–21) mm in length, inner phyllaries
12–15 mm in length................................5. S. amasiana
− Plants not caespitose, glabrous, mature capitula 20–
40 mm in length, inner phyllaries (14–)15–30(–38)
mm in length ................................................................50
50- Achene surface distinctly toothed and winged, pappus
brownish ........................................................31. S. pacis
− Achene surface smooth, pappus cream or dirty white
........................................................................................51
51- Leaves herbaceous, phyllaries glabrous, achene 9–11
mm in length, pappus plumose below and scabrid
above .................................................46. S. tuzgoluensis
− Leaves not as above, inner phyllaries sparsely crisped
pubescent, achene 12–15 mm in length, some pappus
plumose, others scabrid..............................38. S. renzii
Acknowledgments
The authors wish to express their thanks to TÜBİTAK
(109T972) for its financial support, Dr Ahmet Duran for
critical reading and comments, and Mrs Taiadjana Fotuna
for linguistic checking.
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COŞKUNÇELEBİ et al. / Turk J Bot
Appendix. List of examined taxa.
Scorzonera ahmet-duranii: C2 Muğla: Köyceğiz,
Sandras mountain, Topuklu-fire tower, Pinus clearance,
1655 m, 23.06.2010, Makbul & Coşkunçelebi 230 (RUB,
KTUB). Scorzonera bella: C10 Hakkâri: Cilo mountain in
Diz stream, 1730 m, 06.08.1954, Davis & Polunin 23925
(LE). Scorzonera boissieri: B6 Kayseri: Sarız, Yeşilkent,
Binboğa mountains, Tekke rock, mountain steppe, 1879
m, 15.07.2010, Makbul & Coşkunçelebi 250 (RUB, KTUB);
Maraş: Göksun, Berit mountain, above Çavdar, rocky
slopes, 2203 m, 14.07.2010, Makbul & Coşkunçelebi 247
(RUB, KTUB). B6 Kayseri: Sarız, Yalak, Binboğa
mountains, Tekke rocky places, steppe, 2000–2200 m,
09.07.1992, Aytaç & Duman 5157 (HUB); Kayseri: Sarız,
Keklikoluk rock, Işık mountain, Binboğa mountain,
mountain steppe, 2400 m, 12.09.1991, Aytaç 4451 (HUB);
Kayseri: Binboğa mountain, Körkuyu-Sıcak plateau, stony
steppe, 2400–2600 m, 21.07.1992, Aytaç & Duman 5444
(GAZI); Maraş: Göksun, Ericek, Berit mountain, above
Çavdar plateau, 2200–2250 m, 29.06.1992, Aytaç & Duman
5004 (GAZI). Scorzonera coriacea: B3 Isparta:
Şarkikaraağaç, Kızıldağ National Park, 1600–1700 m,
25.06.1994, B.Mutlu 579-928 (HUB); C3 Konya:
Derebucak, Çamlık, Kızıldağ, serpentine places, 1400 m,
12.06.2009, N 37 21 869 E 031 40 501, A. Duran 8349 & B.
Doğan (holo.: KNYA; iso.: GAZI, ANK, HUB, Selçuk
Univ., Herbarium of the Faculty of Education); 1445 m, N
37 21 028-E 031 39 082, 30.05.2005, E.Hamzaoğlu 3707 &
Aksoy (KNYA); Suluin road, serpentine places, 1300 m, N
37 21 299 E 031 36 467, 11.06.2008, A. Duran 8042
(KNYA); Konya: Derebucak, Çamlık road, opposite the
fountain, Pinus clearance, 1401 m, 12.06.2009, Makbul &
Coşkunçelebi 144 (RUB, KTUB); C3 Isparta: Şarkikaraağaç,
Kızıldağ National Park, around the hill, subalpine, alpine,
1599 m, 14.07.2012, Makbul & Coşkunçelebi 354 (RUB,
KTUB). Scorzonera davisii: C10 Hakkari: 30 km from
Hakkari to Başkale, stony slopes, 1430 m, 19.07.2011,
Makbul & Coşkunçelebi 320 (RUB, KTUB); Van: Başkale–
Hakkari road, 20–25 km from Hakkari, Zap gorge,
roadside, slopes, 1533 m, 19.07.2011, Makbul &
Coşkunçelebi 315 (RUB, KTUB). Scorzonera elata: A2 (A)
İstanbul: Kartal, Cevizli, Dragos mountain, hill, Pinus
forest, 28.04.1968, A. Çır. and O. Sut. (ISTF); İstanbul:
Büyük island, 0–100 m, Sibthorp; Bursa: Bursa
surroundings, Sibthorp; A4 Kastamonu: Daday, Juniperus
forest, 700 m, 26.06.1980, Ketenoğlu 792 (ANK); A5
Kastamonu: Tosya, Kavakçeşme, Sint. 1892: 4918b; Tokat:
Zile, Akçakeçeli village, Quercus forest, calcareous stones,
850–950 m, 24.07.1993, Dönmez 3789 (HUB); B1
Balıkesir: Ayvalık, Pınar island, 10 m, 10.05.1996, K.
Akpınar (ISTE); İzmir: İzmir surroundings, Montbret;
İzmir: Bornova–Manisa 15 km, hill 430 m, 08.07.1993, Ö.
Seçmen 4291 (EGE); İzmir: Bayraklı, meadows, 200–300
m 02.06.1933, O. Schwarz (EGE); İzmir: Selçuk–Kuşadası,
Samsun mountain, 24.06.1965, Regel (EGE); Manisa: 20
km S of Manisa, Gediz, 500 m, 26.07.1962, M. & D. Zohary;
B3 Eskişehir: Türkmen mountain, Arban stream, Üçsaray,
P. sylvestris clearance, 16.06.1976, T. Ekim, (ISTE); Konya:
Akşehir, above Atakent, 1080–1200 m, 28.05.1974, G.
Dökmeci and Y. Doğantaş (ISTE); B4 Ankara: Mamak,
Kıbrıs village valley, Cehrelik, grassland, 1300–1350 m,
04.07.2005, Aslan 2419 (GAZI); Ankara: Ayaş Beli, Gazi
Mustafa Kemal Forest, forest clearance, 1100–1200 m,
09.07.1986, Aytaç 2179 (GAZI); B6 Sivas: 12 km N of Sivas,
1600 m, Sorger 60-18-17; B7 Sivas: Divriği, 1000 m, Bornm
1893: 242a; Erzincan: Salihli, 1250–1300 m, Hub.-Mor.
9117; C1 Aydın: Priene residues, 250 m, 25.05.1962, D.
34975 (E); Aydın: Samsun mountain, Regel (EGE 8296);
İzmir: Selçuk, 300 m, Sorger 63-13-4; Muğla: Bodrum, S
of Turgut Reis, W foot of Doru mountain, 16.05.1984, E.
Tuzlacı (ISTE); C2 Aydın: from Aydın to Muğla 72 km, 330
m, Hub.-Mor. 13193; Antalya: Akdağ, 1400 m, Sorger 6722-75; Burdur: 3 km N of Dirmil, 1250–1300 m, Hub.Mor. 9115; Denizli: Denizli–Kaçık Beli road, Karatepe,
Gökkaya plantation area, 750 m, 07.06.1973, A.Baytop and
T.Tuzlacı (ISTE); Denizli: Babadağ, Antalya–Tavas
junction, roadside, 500 m, 20.07.1997, S. Oluk 2656 (EGE);
Muğla: Küçük Kargı, from Fethiye to Köyceğiz, Bozakman
& Fitz 344; Muğla: Ortaca, Tepearası, Eskiköy, Pinus brutia
forest, 60–80 m, 25.05.1991, Vural 5873, (HUB); Muğla:
Köyceğiz, Kavakarası village, maquis, 20 m, 18.05.1991,
Güner 9047 (HUB, GAZI); Muğla: Köyceğiz, Hamitköy,
Domuzdireği hill, maquis, 10–30 m, 21.05.1991, Güner
9169 (HUB); Muğla: Köyceğiz, Ekincik village, Sandallı
hill, 400–600 m, 16.06.1991, Güner 9394 (HUB); Muğla:
Köyceğiz, Beybası village, Süpürgelik hill, maquis, 150 m,
21.04.1991, Güner 8994, (HUB); Muğla: Dalaman, from
Dalaman to Köyceğiz 10 km before, P. brutia forest,
roadside, 152 m, 25.05.2010, Makbul & Coşkunçelebi 204
(RUB, KTUB); C3 Antalya: Tahtalı mountain, Tekirova, 30
m, Hub.-Mor. 9983; Antalya: Adrasan, Çavuş village, Sazak
road, 15 m, 15.05.2009, Eustace Ann 118 (E); Antalya:
Akseki–Güzelsu, Serebil area, Pinus brutia forest, 1100 m,
08.07.1996, Duran 4136, (GAZI); C4 Antalya: Alanya,
Çetik 124; Antalya: Alanya, Mahmutlar, Kuşkayası, Pinus
nigra forest, 1371 m, 04.07.2010, Makbul & Coşkunçelebi
236 (RUB, KTUB); C6 Maraş: 3 km S of Süleymanlı,
shrubs, 900 m, 16.06.1981, Yıldız 2811 (HUB!). Scorzonera
hieraciifolia: B4 Konya: Gölyazı plateau, salty places, 912
m, 11.06.2009, Makbul & Coşkunçelebi 138 (RUB, KTUB);
B4 Aksaray: Tuz lake, Aksaray, Eskil-lake, swamp, Juncus
community, 940 m, 02.06.1998, Aydoğdu (GAZI); Konya:
Cihanbeyli, 800 m, Hub.-Mor. 14690; B5 Kayseri: İncesuDeveli, 1200 m, salty swamp, 29.08.1957, D. 32766 (Typus)
(E); Kayseri: İncesu-Develi, 1200 m, D. 32766; Kayseri:
Sultan reeds, Yahyalı, Ovaçiftliği village, Kuş museum,
1
COŞKUNÇELEBİ et al. / Turk J Bot
protected area, 1071 m, 10.07.1993, Yıldırımlı 16171
(HUB); Kayseri: Yahyalı, Sultan reeds, Tuzla, 03.07.1982,
Sümbül 1475 (HUB); Kayseri: Yahyalı-Sultan reeds, Tuzla,
03.07.1982, Demirkuş 1913 (HUB); Niğde: Aksaray,
Birand & M.Zohary 2819; B6 Sivas: Tödürge lake, ZaraHafik, 1350 m, Hub.-Mor. 13204; B6 Sivas: Tödürge lake, S
of Zara-Hafik, salty places, 1301 m, 06.06.2009, Makbul &
Coşkunçelebi 127 (RUB, KTUB). Scorzonera inaequiscapa:
A7 Giresun: 10–15 km from Alucra to Şiran, Hacıhasan
village, grassy and rocky area, 1670 m, 13.06.2005, Makbul
79 (RUB, KTUB); Giresun: Alucra–Şiran, Hacıhasan
village, grassland, sandy and rocky area, 1661 m,
27.06.2011, Makbul & Coşkunçelebi, 298 (RUB, KTUB);
A8 Bayburt: 2.5 km from Bayburt to İspir, roadside, 1506
m, 30.05.2005, Makbul 68 (RUB, KTUB); B5 Yozgat:
Akdağmadeni forest station, 06.06.1960, Curtis 177; B7
Tunceli: Ovacık, Karagöl valley, Munzur mountain, 1400
m, 05.05.1979, Yıldırımlı 1347 (HUB). Scorzonera
karabelensis: C2 Muğla: Fethiye, Fethiye–Korkuteli road,
clearance of Pinus forest, rocky chalky slopes, 1114 m,
22.05.2012, Makbul & Coşkunçelebi 346 (RUB, KTUB).
Scorzonera longiana: C4 Antalya: Gazipaşa, Çayıryakası
plateau, 1700 m, 14.07.1983, Sümbül 2290 (HUB); Antalya:
Gazipaşa, from Çobanlar village, plateau to Oyuklu
plateau, 1800–2000 m, 11.07.1983, Sümbül 2239 (types
HUB, E). Scorzonera longiana: C3 Antalya: Gazipaşa,
Çayıryaka plateau, stony meadows, 1727 m, 25.07.2010,
Makbul & Coşkunçelebi 255 (RUB, KTUB). Scorzonera
pacis: C6 Hatay: Antakya, Karaali, maquis, 410 m,
10.04.2011, S. Yıldız 551 & S. Kayıkçı (holo.: GAZI, iso.:
HUB, MKU). Scorzonera phaeopappa: C3 Antalya:
Akseki, Yaylacık hill, 1900 m, Quezel et al.; C5 Adana:
Adana–Ceyhan, 6 km E of Ceyhan stream, Coode & Jones
3571; Adana: Mısır–Ceyhan, 25 m, 14.04.1956, D. 26069
(ANK); Adana: Mısır, 11.04.1934, Balls 717 (ANK); Adana:
8 km from Adana to Kozan, calcareous soils, 150 m,
12.04.1957, D. 26628 (E); Niğde: Ala mountain, Çukurbağ–
Narpız, 1900–2050 m, Parry 72; C6 Gaziantep: Dülük
Baba, 7 km N of Gaziantep, 1100 m, 12.05.1957, D. 27879
(ANK, E); Gaziantep: Dülük Baba, Gaziantep, 1250 m,
Haradj. 1217; Gaziantep: Dülük Baba, picnic area, Pinus
forest, 1135 m, 15.05.2010, Makbul & Coşkunçelebi 174
(RUB, KTUB); Hatay: Kırıkhan, Antalya–İskenderun, 5
km from Kırıkhan junction, roadside, 385 m, 16.05.2010,
Makbul & Coşkunçelebi 177 (RUB, KTUB); Maraş: Ahır
mountain, Yalnız Ardıç backs, 1300–1500 m, 22.05.1992,
Aytaç & Duman 4668 (HUB); Maraş: Engizek mountain,
Aksu, Juniperus excelsa clearance, 1000–1200 m,
24.05.1987, Duman 2919 (GAZI); Maraş: Ahır mountain,
Akdere, 1300–1600 m, 03.05.1991, Aytaç 3564 (GAZI); C7
Urfa: Ceylanpınar, Sorkah, 470–490 m, 14.04.1995,
Adıgüzel 1895 (GAZI); C8 Batman: Bismil, 5 km from
steppe area to hill, 610 m, 25.04.2009, K 37 81 06-D 40 72
2
53, Aslan et al. 3210 (GAZI); Mardin: 24 km from
Diyarbakır to Mardin, 1000 m, D. 28735; Urfa: Ceylanpınar,
Hausskn. Scorzonera pisidica: C2 Burdur: Tefenni,
Quercus coccifera, 3 km from Altınyayla, 1200–1270 m,
27.06.1948, Hub.-Mor. 8485 & Reese (holo Hb. Hub. Mor.);
Burdur: 6 km S of Dirmil, 1600–1650 m, Hub.-Mor. 8486
& Reese; Burdur: Gölhisar-Dirmil, 1270 m, 17.06.1981,
Nydegger 16244 (tip) (E); Burdur: Yeşilova, S of Salda lake,
P. nigra and Quercus clearance, 1170–1200 m, 11.07.1993,
Duman 5080 (GAZI); C2 Denizli: Honaz, roadside, stony
field, Juniperus clearance, 1743 m, 13.07.2012, Makbul &
Coşkunçelebi 353 (RUB, KTUB); Muğla: Köyceğiz,
Sandras mountain, climbing from Beyağaç to Sandras
mountain, Yumaklı area, Pinus sp. forest, 1282 m,
23.06.2010, Makbul & Coşkunçelebi 229 (RUB, KTUB);
Muğla: Köyceğiz, Sandras mountain, Topuklu–Yangın fire
tower, Pinus sp. forest, 1655 m, 23.06.2010, Makbul &
Coşkunçelebi 231 (RUB, KTUB); Muğla: Beyağaç–Sandras
mountain, P. nigra forest, 1320 m, 16.06.2001, Varol 3895
(GAZI). Scorzonera pygmaea: A2 Bursa: Uludağ, Juniperus
clearance, 2159 m, 24.07.2010, Makbul & Coşkunçelebi
271 (RUB, KTUB); A4 Kastamonu: Ilgaz mountain,
around TV transmitter, alpine, 2055 m, 02.08.2009,
Makbul 165 (RUB, KTUB); A5 Kastamonu: Ilgaz
mountain, back hills of the TV transmitter, steppe, 1998
m, 20.08.2011, Makbul & Coşkunçelebi 339 (RUB, KTUB).
A2(A) Bursa: Uludağ, 19.08.1850, Clementi; 2400 m,
Sorger 68-54-23; Bursa: Uludağ, Karagöl road, 24.08.1971,
A. & T. Baytop (ISTE); Bursa: Uludağ, alpine, 13.08.1953,
A. & T. Baytop (ISTE); 29.08.1950, A. & T. Baytop (ISTE);
A2 Bursa: Uludağ, 2380–2580 m, 13.09.1949, D. 1484614862a (E); Bursa: Uludağ, 29.07.1968, Pamukçuoğlu
(HUB); Bursa: Uludağ, NW of hill, calcareous stones,
2300–2500 m, 05.08.1993, Y. Gemici 8284 (EGE); Bursa:
Uludağ, around the hill, stream bed, 1950 m, 27.07.1984,
Ö. Seçmen 11 (EGE); Karabük: peak of Keltepe, 1950 m,
12.07.1984, Demirörs 1263 (ANK); Kastamonu: Ilgaz
mountain, limy stones, 2200 m, 28.07.1962, D. 2157738376 (E); Kastamonu: Ilgaz mountain, Pinus forest, 2000
m, 19.06.1969, Darrah 28 (E); Kastamonu: Ilgaz mountain,
K. Hacet mountain, S slopes of alpine, limy, 2400 m,
29.02.1982, Akman et al. 12178 (ANK, EGE); A4
Zonguldak: Karabük-Keltepe, 1950 m, calcareous hills,
03.08.1962, D. 39814 (E); B2 Kütahya: Gediz, Şaphane
mountain, calcareous stones, 1700–1900 m, 17.06.1993,
Y.Gemici 7695 (EGE); C3 Isparta: Dedegöl Mountain,
1600–2400 m, Sorger 70-46-24. Scorzonera rigida: B7
Erzincan: above Ahmediye, NW slopes, alpine, stony
places, 2306 m, 11.07.2009, Makbul 156 (RUB, KTUB); C4
Konya: Ereğli, Kayasaray village, Düğünlük stream,
calcareous stones, 1910 m, 05.07.2010, Makbul 240 (RUB,
KTUB). B7 Erzincan: Sipikör mountain, 1990 m, Hub.Mor. 13194; B9 Bitlis/Van: 10 km SE of Pelli, calcareous
COŞKUNÇELEBİ et al. / Turk J Bot
slopes, 2600 m, 08.07.1954, D. 22544 (ANK, E); Van:
Başkale, İspiriz mountain, stony slopes, 3200 m, 31.07.1954,
D. 23763 (ANK, E); Van: 8 km from Van to Erçek, stony
slopes, 2100 m, 06.06.1966, D. 44424 (ISTO, E); Van:
Çatak, Kavussahap mountain, 3100 m, 23.07.1954, D.
23122 (E); Van: Gevaş, Artos mountain, rocky slopes, 2134
m, 14.07.1954, D. 22681 (E); Van: 36 km from Başkale to
Hoşap, Güzel Dere pass, rocky-steppe slopes, 2750 m,
03.07.1966, D. 45989 (E); Van: Başkale, Işıklı village,
13.06.1984, T. Baytop (ISTE); Van: Erek mountain,
07.07.1988, Özçelik & Ay (EGE); Van: Erek mountain,
3100 m, Kronenburg 130; B10 Ağrı: 3 km E of Doğubeyazıt,
calcareous stones, 1750 m, 31.05.1966, D. 43945 (ISTO, E);
C5 Adana: Gülek, 2000 m, Sorger 62-71-23; C5 Mersin:
Yıldız hill, SE slopes, 2700–3000 m, 17.07.1989, Y. Gemici
4860 (EGE); C5 Niğde/İçel: Bolkar mountain, W of
Mededsiz hill, stony slopes, 2500–2600 m, 26.07.1984,
Strid et al.. 24036 (EGE); C6 Maraş: Berit mountain, 1700–
2500 m, Hausskn.; C9 Van: Harefta mountain, Başkale–
Koçanis, 2700–2800 m, Nábělek 3737; C10 Hakkari: Cilo
mountain, 10 km W of Cilo hill, 3700 m, 09.08.1954, D.
24216 (ANK). Scorzonera sandrasica: C2 Muğla: Köyceğiz,
Sandras mountain, Beşparmak hill, around fire tower,
steppe areas, 1970 m, 13.04.2004, Varol 5634 (GAZI);
Muğla: Sandras mountain, W of the summit, 1970 m,
13.04.2004, N 37 04 E 028 50, Hartvig (izo. EGE); C2
Muğla: Köyceğiz, Beşparmak hill, around fire tower, rocky
slopes, forest boundaries, 2025 m, 23.06.2010, Makbul &
Coşkunçelebi 232 (RUB, KTUB). Scorzonera sublanata:
A1 Çanakkale: Eceabat, Yalova village, Cistus shrubs, hill,
110 m, 02.06.2011, Makbul & Coşkunçelebi 288 (RUB,
KTUB); A1(E) Çanakkale: S of Eceabat, 300 m, Sorger 637-14; Çanakkale: Gökçeada, Tuzburnu–Hacıahmet, 70 m,
15.04.1976, Ö. Seçmen 1575-1588 (EGE); 60 m, 05.05.1975,
Ö. Seşmen 428 (EGE); A1(A) Çanakkale: Kemerdere, Sint.
1883: 325; A4 Ankara: Çubuk, Ovacık, Saraycık village,
Atcameydan hill, shrubby slopes, 1250–1380 m,
20.05.1992, Dündar 495 (GAZI); B1 Balıkesir: Ayvalık,
Cunda island, above Patriçe, 10 m, 07.05.1996, K.Alpınar
(ISTE); Balıkesir: Ayvalık, Armutçu–Sakarya, 20.04.1997,
K.Alpınar (ISTE); Balıkesir: Ayvalık, Dolap island–
Mitralyöz, 5 m, 23.05.1997, K.Alpınar (ISTE); Balıkesir:
Ayvalık, Hakkıbey S of island, 15.04.1998, K.Alpınar
(ISTE); İzmir: Gümüldür, W of Değirmendere, 15.04.1969,
Fitz & Spitz.; İzmir: Karaburun, Balıkova–Gerence,
05.04.1980, L.Bekat 240 (EGE); İzmir: Değirmendere–
Gümüldür, 15.04.1969, K.Fitz (EGE); Manisa: Spil
mountain, rocky and grassy, Juniperus clearance, 1224 m,
22.05.2011, Makbul & Coşkunçelebi 280 (RUB, KTUB);
Manisa: Spil mountain, steppe, 1196 m, 21.06.2010,
Makbul & Coşkunçelebi 228 (RUB, KTUB); B2 Uşak: Uşak
surroundings, 900 m, Bal.; B4 Ankara: Ankara, 1100 m,
Balls 1933: 221; Ankara: Şereflikoçhisar–Aksaray road, 10
km from Şereflikoçhisar, steppe, 935 m, 24.05.2010, N 38
49 760 E 033 35 921, Makbul & Coşkunçelebi 199 (RUB,
KTUB); C2 Aydın; below Karacasu, 400–500 m, maquis,
23.04.1965, D 41633 (E); Aydın: Aydın–Muğla, Pinus
clearance, 458 m, 19.05.2011, Makbul & Coşkunçelebi 275
(RUB, KTUB); Muğla: Yılanlı mountain, rocky area, 1369
m, 19.05.2011, Makbul & Coşkunçelebi 276 (RUB, KTUB);
Denizli: Pamukkale, 07.05.1969, Fitz & Spitz.; Denizli:
Babadağ, Karacasu, above Yenice, P. brutia forest, 200 m,
24.04.1998, S.Oluk (EGE); Antalya: Elmalı, Bourgeau 169;
C3 Antalya: Bük forest, Kayran, Bozakman & Fitz 149;
Antalya: Kumluca, W of Adrasan village, serpentine
slopes, P. brutia forest, 10–50 m, 29.04.1980, Peşmen 4881
(HUB); Antalya: Kumluca, Adrasan village, Musa
mountain, Kızıl area, serpentine slopes, P. brutia forest,
0–150 m, 23.03.1979, Peşmen 4243 (HUB); Antalya:
Kemer, Olimpos, 50 m, 23.03.1979, Peşmen 4229 (HUB).
Scorzonera tomentosa: A4 Ankara: Kalecik, 1 km W of the
village, serpentine, 12.07.1998, Adıgüzel 1899 (E); Ankara:
Çubuk, Karagöl, Yaşmağın hill, 1500 m, 05.08.1974, Erik
683 (HUB); Çankırı: Çankırı, 800 m, Bornm. 1929: 14326;
A5 Amasya: Direkli village, Çetinkaya, 1550 m, 03.10.1976,
K.Akpınar (ISTE); Kastamonu: Tosya, Sint. 1892: 4721; A6
Sivas: Yıldız mountain, Cırcır, steppe, 1400 m, 14.08.1967,
Tobey 2312 (E); A7 Gümüşhane: Kop village, rocky slopes,
1300 m, 19.06.1967, Tobey 2043 (E); A8 Artvin: Artvin,
Barevan, Woronow 6153; Bayburt: Kop mountain,
roadside, 2148 m, 05.07.2007, Makbul 10 (RUB, KTUB);
Bayburt: Bayburt–Aşkale, Kop pass, 2350 m, 16.08.1984,
Leblebici (EGE); Erzurum: 4 km W of Kuzgun village,
rocky slopes, 2250–2350 m, 19.07.1977, Tatlı 5857 (HUB);
Gümüşhane: near Bayburt, Bourgeau 147; A9 Kars: E of
Kağızman mountain, 17 km from Akçay to Cumaçay, 2070
m, 17.07.1966, D. 46720 (ISTO, E); Kars: Demirkapı,
Woronow 12686; B2 Uşak: Bulkas mountain, 1250 m, Bal.;
B3 Konya: Sultan mountain, 28.06.1953, Birand & M.
Zohary; B4 Ankara: near Ankara, Liston; Ankara: Mamak,
Kıbrıs village valley, stony slopes, 1000–1100 m, 01.07.2004,
Aslan 1592 (GAZI); Ankara: Beynam, 400 m, 05.09.1949,
D. 13101; B5 Yozgat: Yıldızlı–Akdağmadeni, 10 km to
Akdağmadeni, stony area, 1113 m, 06.06.2010, Makbul &
Coşkunçelebi 211 (RUB, KTUB); B6 Adana: Binboğa
mountain, Yalak, 2000 m, 14.07.1952, D. 19938 (ANK);
Kayseri: Sarız, Yalak, Binboğa mountain, above Yalak,
rocky field, 1700–2000 m, 08.07.1992, Aytaç & Duman
5242 (GAZI); Kayseri: Akkışla, above Ganişeyk, Hınzır
mountain, 1900 m, 20.08.1979, Çelik 771 (HUB); Kayseri,
Sarız, Yalak, Binboğa mountain, stream side, stony field,
1500–1700 m, 04.08.1991, Aytaç 4349 (GAZI); Kayseri:
Pınarbaşı, Kaynar, S slopes of Hınzır mountain, above
Kara Ahmed, slopes, 2000 m, 18.07.1975, Çelik 531
(ANK); Kayseri: Sarız, Yeşilkent, Binboğa mountain,
climbing to Tekke stone, calcareous stones, 1734 m,
3
COŞKUNÇELEBİ et al. / Turk J Bot
15.07.2010, Makbul & Coşkunçelebi 249 (RUB, KTUB);
Malatya: Doğanşehir, Eskiköy, Çobanderesi-İhtiyarcave,
Juniperus excelsa–Quercus forest, 1600–1800 m,
25.07.1971, Peşmen 2635 (HUB); Malatya: Doğanşehir,
Dedeyazı village, Keşiş mountain, alpine steppe, 1800–
2200 m, 28.07.1971, Peşmen 2580, (HUB); Malatya:
Akçadağ, Kürecik, Aksüt village, 1443 m, 18.07.2005, Y.
Yeşil (ISTE); Maraş: Göksun, Kınıkköz village, Gözpınarı
mountain, 1700 m, 22.08.1977, Yıldız 1532 (HUB); Maraş:
Göksun–Değirmendere, Koruyaz mountain–Meryemçil
protected area, 1750–2000 m, 05.07.1993, Ekici 1577
(GAZI); Maraş: Göksun, Binboğa mountain pass, 1500 m,
14.07.1952 (E); Sivas: Ak mountain, near Sivas, 2200 m,
Siehe 1911: 308; Sivas: Şarkışla, 1500 m, D. 32720 (E);
Sivas: Hekimhan–Kangal, hillside, 1800 m, 02.09.1954, D.
24844 (E); Sivas: 8 km from Sivas to Ulaş, 1300–1450 m,
06.08.1982, D. 68628 (HUB); Sivas: Hafik, Koşutdere–
Bakışecik, stony places, 1250 m, 30.06.1991, Aydoğdu
3245 (GAZI); Sivas: Şarkışla, Evcidere Kale–Alaman,
calcareous stones, 1400–1500 m, 19.07.1979, Ekim &
Düzenli 4022 (ANK); Sivas: Ulaş, Tecer village, Tecer
mountain, 1600 m, 21.07.1978, A.Baytop et al. (ISTE);
Sivas: Çelebiler, Okatan farmer sulfated soils, 1500 m,
03.08.1972, Y.Okatan (ISTF); Sivas/Tokat: Çamlıbel pass,
1650 m, 21.07.1978, A.Baytop et al. (ISTE); B7 Elazığ: Kop
mountain, Sarık village, steppe, 1700 m, 18.07.1984, Evren
2108 (GAZI); Erzincan: Kemaliye (Eğin), Sırakonaklar
village, calcareous stones, 1600 m, 10.07.1982, Tanker
10516 (ANK); Erzincan: Sipikör, Sint. 1890: 3309;
Erzincan: Keşiş mountain, volcanic slopes, 2600 m,
28.07.1957, D. 31759 (E); Erzincan: Erzincan–Refahiye, 25
km E of Refahiye, near Sakallutan pass, serpentine area,
1960–2020 m, 15.08.2001, N 39 52 146 E 039 04 054,
Adıgüzel 4156 (GAZI, E); Erzurum: W of Erzurum,
Erzurum–Kandilli, Ağaver–Kandilli, Tapi stream, limy
slopes, 1960 m, 17.07.1969, O.Özbay (ISTF); Malatya:
Kırlangıç village, Eğri Dere, back of the cave, 1500 m,
15.07.1993, N.Gören (ISTE); Malatya: Doğanşehir,
Eskiköy, above İhtiyar cave, alpine steppe, 1900–2000 m,
14.07.1971, Peşmen 2331 (EGE); Tunceli: Ovacık, Munzur
mountain, Karagöl valley, 1500 m, 09.09.1979, Yıldırımlı
2463 (HUB); Tunceli: Munzur mountain, above Ovacık,
Aksu stream, stony places, 1800 m, 21.07.1957, D. 31470
(E); 2100 m, 19.07.1957, D. 31313 (E); Tunceli: N pass of
Pülümür, near Küçü Dere, rocky slopes, 03.08.1974, Rix
2388 (E); Tunceli/Erzincan: Pülümür pass, grassland, 1950
m, 23.07.197, F. Holtz (EGE); B8 Erzurum: Palandöken
mountain, Kırkdeğirmenler, 1950–2100 m, 22.07.1984,
Demirkuş 2028 (HUB); Erzurum: Tercan–Selepur, 1400
m, 11.07.1957, D. 30954 (E); Erzurum: 66 km from Hınıs
to Erzurum, Aras pass, 1650 m, 12.07.1966, D. 46437
(ISTO, E); C3 Antalya: Gazipaşa–Çayıryaka plateau road,
rocky-stony places, 1714 m, 25.07.2010, Makbul &
Coşkunçelebi 254 (RUB, KTUB); Konya: Beyşehir,
4
Kurucuova, Radar-Karagöl, calcareous stones, alpine,
2000–2500 m, 24.07.1975, Güner 2290 (HUB); Konya:
Kuru tepe, 1400 m, Sorger 66-44-105; C4 İçel: Anamur,
Anamur–Kazancı road, Kızıl Alan, 1300 m, 24.06.1984,
Sümbül 3103 (HUB); Konya: 19 km from Konya to
Beyşehir, 1210 m, Hub.-Mor. 8694; C5 Konya: Ereğli,
Aydos mountain, Berendi, Boğaz, calcareous stones, 1600
m, 28.06.1976, Erik 1705 (HUB); C6 Osmaniye: Zorkun,
Amanos mountain, Keldaz Hill, calcareous stones, 2020 m,
05.07.2010, Makbul & Coşkunçelebi 241 (RUB, KTUB);
C7 Malatya: 74 km SW of Malatya, 1550 m, Sorger 71-4512. Scorzonera tuzgoluensis: B4 Konya: Cihanbeyli,
Gölyazı–Tuzgölü, 908 m, 11.06.2009, N 38 32 552 E 033 21
188, A. Duran 8368 (holo.: KNYA, iso.: GAZI, ANK, HUB,
MR); 12.07.2006, A. Duran 7241 (KNYA, MR); 29.06.2009,
B. Doğan 2100 (KNYA, GAZI, MR); Konya: Cihanbeyli,
climbing to Gölyazı plateau, Dumanağlı plateau, salty
swamp, 908 m, 06.07.2011, Makbul & Coşkunçelebi 313
(RUB, KTUB). Scorzonera ulrichii: C4 Antalya: Alanya,
Çökele–Gökbel, calcareous area, near Pinus forest, 1425
m, 05.07.2003, Göktürk 5107 (HUB, GAZI); Antalya:
Alanya, Mahmutlar–Hadim road, c. 35 km NE of
Mahmutlar and 15 km S Çayarası, 1 km N Elmalısu,
gravely, rocky slopes with Pinus nigra var. caraminaca
forest clearance, W-exp., limestone, 1250 m, 11.06.2002,
Robert Ulrich (Tubingen) 2/12 (holo.:B, iso.: E, ISTE); C4
Antalya: Alanya, Mahmutlar, Elmalısu–Hadim road, after
Kuşkaya, calcareous rocks, under Pinus nigra, 1336 m,
05.07.2010, Makbul & Coşkunçelebi 237 (RUB, KTUB);
Antalya: Alanya, Mahmutlar–Hadim road, Elmalısu,
junction of Gökbel plateau road, calcareous rocks, under
Pinus nigra, 1336 m, 05.07.2010, Makbul & Coşkunçelebi
238 (RUB, KTUB). Scorzonera veratrifolia: B9 Bitlis:
Kambos mountain, above Hurmuz, 1830 m, 31.06.1954, D.
23494 (E)!; Van: Pelli mountain, 54 km SW of Van 2500 m,
09.07.1971, Edmondson 618 (E)!; Van: Van lake, 20 km SE
of Tatvan, 1960 m, Frodin 1939, 1971 and 1972; C9
Hakkari: between Beytuşşebap and Uludere, W of Tanin
mountain, 2500 m, 25.07.1974, Rix 2338 (E)!; Hakkari:
Cilo mountain, Diz stream, 1740 m, 06.08.1954, D. 23926
(E)!; Van: Çatak, Dalbastı village, 1350–1600 m, 20.07.2002,
Bani 1259 (GAZI)!; C10 Hakkari: Sat mountain, 2000–
2100 m, 30.06.1966, D. 45812 (ISTO, E)!; Hakkari Cilo
mountain, 2500 m, 25.08.2011, MF 27719!; Hakkari:
Koçanis, 2500 m, 21.07.2011, MF 27566!; Hakkari:
Esendere, Mor mountain, 2700 m, 24.07.2011, MF 27194!;
Hakkari: Diz stream, 2200 m, 23.07.2011, MF 27168!.
Scorzonera yildirimlii: C6 Osmaniye: Amanos mountain,
Zorkun plateau, Keldaz hill, 1950 m, 05.07.2001, N 36 58
95 E 036 24 22, Duran 5765-6824 (iso.: HUB, GAZI; para.:
E); Osmaniye: Zorkun, Amanos mountain, Keldaz hill,
Halep Gösteren, calcareous rocky hills, 2073 m, 05.07.2010,
Makbul & Coşkunçelebi 243 (RUB, KTUB).