Turkish Journal of Botany
http://journals.tubitak.gov.tr/botany/
Research Article
Turk J Bot
(2016) 40: 87-96
© TÜBİTAK
doi:10.3906/bot-1407-71
New species and a synonym of the genus Hesperis (Brassicaceae) from Turkey
Ahmet DURAN, Özlem ÇETİN*
Department of Biology, Faculty of Science, Selçuk University, Selçuklu, Konya, Turkey
Received: 18.07.2014
Accepted/Published Online: 12.04.2015
Final Version: 01.01.2016
Abstract: Hesperis tosyaensis A.Duran, a new species from northern Anatolia, is described and illustrated. It grows under open Quercus
forest. The new species is distinct from the closely allied H. ozcelikii A.Duran by the following features: stem erect, 45–90 cm tall,
hispid, hairs 2–2.5 mm, and fruit glabrous. Genetic differentiation between closely related Hesperis L. taxa and Matthiola W.T.Aiton was
examined by molecular analyses of the ISSR. The chromosome number for Hesperis tosyaensis is 2n = 14. Information is also presented
on its ecology and conservation status. In addition, Hesperis aspera E.Fourn. is treated as a synonym of Hesperis bicuspidata (Willd.)
Poir.
Key words: Hesperis, new species, ISSR, Cruciferae, systematics
1. Introduction
Hesperis L. (Brassicaceae) is represented by many taxa at
the junctions of the Irano-Turanian, Mediterranean, and
Euro-Siberian phytogeographic regions; these regions
meet in Anatolia. Towards the outer boundaries of each
phytogeographic region, Hesperis is represented by fewer
taxa, which is also indicated by the number of Hesperis taxa
in various floras. There are 14 species in Europe, 11 species
in Iran, 9 species in Romania, 5 species in Iraq, 3 species in
Italy, and 1 species in Palestine (Săvulescu, 1955; Ball, 1964;
Zohary, 1966; Dvořák, 1968, 1980; Pignatti, 1982; Duran
and Hamzaoğlu, 2004; Duran and Ocak, 2005; Duran, 2005;
Parolly and Tan, 2006; Duran, 2008, 2009).
The genus Hesperis is represented by 31 species in the
flora of Turkey (Cullen, 1965; Davis et al., 1988). Five new
species have been described from Turkey and 10 species
are treated as synonymous (Duran and Ocak, 2005; Parolly
and Tan, 2006; Duran, 2008, 2009; Duran et al., 2011).
According to the latest study, Hesperis is represented by 27
species recorded from Turkey (Duran, 2012).
The genus Hesperis in Turkey has been studied in terms
of morphology, floristic features, palynology, molecular
characteristics, anatomy, and chemotaxonomy (Duran,
2005; Duran and Ocak, 2005; Duran, 2008; Ünal et al.,
2008; Yücel et al., 2008; Aras et al., 2009; Büyükkartal et al.,
2009; Duran, 2009; Pınar et al., 2009; Kırımer et al., 2010).
Despite the fact that many studies have been performed
*Correspondence: [email protected]
on the infrageneric and infraspecific classification of the
genus Hesperis, ecological and geographic variations of
some Hesperis taxa cause some confusion in taxonomy
and nomenclature, especially among Hesperis matronalis
L., H. bicuspidata (Willd.) Poir., H. pendula DC., and
certain other taxa. Hesperis bicuspidata is localized in the
Irano-Turanian phytogeographical region and the species
has a widespread distribution in Turkey. H. bicuspidata
grows in different habitats with different features. Hesperis
bicuspidata specimens were collected from different
localities in Turkey.
Hesperis aspera, which is endemic to Turkey and
is known only from Tosya (Kastamonu Province), was
described as a new species by Fournuer (1866). Our
examinations of type specimens and herbarium specimens
of H. aspera have revealed that it is a synonym of H.
bicuspidata. Based on our study, H. aspera is treated as a
synonym of H. bicuspidata.
Some interesting Hesperis specimens were collected
from the Tosya district (Kastamonu). Herbarium studies
and field observations showed that our specimen is a
new taxon. Diagnostic morphological characters from
closely similar Hesperis species are discussed in this study.
Ecology, conservation status, and biogeography of the new
species are presented. Genetic differentiation of the new
species and related species were analyzed using NTSYSpc
2.1 software and dendrograms were obtained.
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DURAN and ÇETİN / Turk J Bot
2. Materials and methods
2.1. Morphology
We followed the descriptions of taxa according to the
Flora of Turkey and the eastern Aegean Islands and
other floras (Halácsy, 1900; Hayek, 1927; Busch, 1939;
Săvulescu, 1955; Tzvelev, 1959; Ball, 1964; Cullen, 1965;
Zohary, 1966; Dvorák, 1968; Dvorák, 1980; Pignatti, 1982;
Davis et al., 1988; Tan and Iatrou, 2001) were examined.
The specimens present in the AEF, ANK, BM, E, EGE, G,
GAZI, HUB, ISTF, K, KNYA, P, and WU herbaria were
examined. All measurements were made on 10 randomly
selected specimens. Measurements were made of plant
parts using a ruler and an ocular micrometer under a
microscope. Localities of studied taxa are given in the
Appendix.
2.2. Karyology
Karyological observations were carried out on root
tips obtained from germinating seeds. Root tips were
pretreated for 16 h in α-monobromonaphthalene at 4 °C
and fixed in Carnoy solution (3:1 absolute ethanol and
glacial acetic acid), and then the root tips were hydrolyzed
with 1 N HCl at room temperature and stained with 2%
aceto-orcein. Stained root tips were flattened in a drop
of 45% acetic acid, and permanent slides were made by
mounting in Depex.
2.3. Molecular analyses
Total DNA was obtained from 50–75 mg of dried leaf
tissue from 11 different individuals. DNAs were extracted
using phenol-chloroform (Sambrook et al., 1989) after
concentrations were determined by NanoDrop. Sample
DNAs were diluted to 25 ng/µL. Stock DNAs were kept
at –86 °C. ISSR primers were used for PCR amplification;
the characteristics of the primers used are given in Table
1. Each reaction contained 25 μL. PCR reactions were
performed in a mix containing 2.5 μL of PCR buffer, 3 μL
of 25 mM MgCl2, 1 μL of primer, 0.5 μL of dNTP mix, 0.3
μL of Taq DNA polymerase, 4 μL of each diluted DNA, and
13.7 units of PCR water. After a predenaturation step of 3
min at 94 °C, amplification reactions were cycled 40 times
at 94 °C for 1 min, at annealing temperature (Table 1) for 1
min, and at 72 °C for 1 min in an Eppendorf Mastercycler
gradient thermocycler. A final extension was performed
for 10 min at 72 °C. Amplified fragments were visualized
under a UV transilluminator and photographed using
a gel documentation system (Vilbert Lourmat, Infinity
model). All amplified fragments were treated as dominant
genetic markers. Each DNA band generated was visually
scored as an independent character or locus. Analyses
were performed with the NTSYS-pc 2.1 software package
(Rohlf, 2001). The dendrogram was constructed using SM
coefficient and UPGMA clustering methods.
3. Results
Hesperis bicuspidata (Willd.) Poir., Encycl. Suppl. 3: 195
(1813)
Sect. Hesperis
Type: Tournefort, no. 16 (B, P! KNYA photo.!, GAZI
photo!). Described from Turkey.
Synonyms ≡ Cheiranthus bicuspidatus Willd., Sp. Pl., 3,
1: 519 (1800). = Hesperis kotschyana Fenzl, Pug. PI. Nov.
Syr., 13 (1842); H. violacea Boiss., Diagn. PI. Orient., ser.
1, 5: 80 (1844); H. reuteriana Boiss. & A.Huet, Diagn. PI.
Orient., ser. 2, 5: 20 (1856); H. unguicularis Boiss., Diagn.
PI. Orient., ser. 2, 5: 21 (1856); H. aspera E. Fourn., Bull.
Soc. Bot. Fr. 13: 352 (1866) syn. nov.; H. tauricola Kotschy
& Boiss., Fl. Orient. 1:232 (1867); H. sintenisii Hausskn.
& Bornm., Repert. Spec. Nov. Regni Veg. Beih. 89(1): 38
(1936); H. stellata Dvorak, Oesterr. Bot. Z. 112: 610 (1965).
Hesperis tosyaensis A.Duran sp. nov. (Figures 1–6)
Table 1. The characteristics of the ISSR primers.
Primer
Sequences
Tm (°C)
Length (bp)
GC ratio (%)
Total bands
M1
5ʹ-AGC AGC AGC AGC AGC AGC G-3ʹ
63.10
19
68.4
5
M2
5ʹ-ACC ACC ACC ACC ACC ACC G-3ʹ
63.10
19
68.4
9
M3
5ʹ-AGC AGC AGC AGC AGC AGC C-3ʹ
63.10
19
68.4
9
M5
5ʹ-GAG AGA GAG AGA GAG AGA C-3ʹ
56.70
19
52.6
16
M7
5ʹ-AGA GAG AGA GAG AGA GAG C-3ʹ
56.70
19
52.6
15
M9
5ʹ-ACA CAC ACA CAC ACA CCG-3ʹ
56.00
18
55.6
8
Diplo1
5ʹ-CAC ACA CAC ACA CAC AG-3ʹ
57.19
17
52.9
7
Cucurbita
5ʹ-GAG CAA CAA CAA CAA CAA -3ʹ
53.07
18
33.3
25
Cucurbita2
5ʹ-GAG AGA GAG AGA GAG AGA T-3ʹ
54.78
17
52.9
10
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DURAN and ÇETİN / Turk J Bot
Figure 1. Hesperis tosyaensis: A- general view, B- mature fruit. Scale bar = 2 cm.
Subgen. Hesperis Sect. Hesperis
Type: Turkey. A5 Kastamonu: between Tosya-Sekiler
village, 7 km, 1000 m, under an open Quercus forest,
40°55.62′N, 34°02.06′E, 12.5.2001, A.Duran 5657 &
Hamzaoğlu (holotype: KNYA, isotypes: GAZI, ANK,
ISTE, ISTF, E, K).
Diagnosis: Hesperis tosyaensis differs from H. ozcelikii
by its erect stem, 45–90 cm (not ascending to erect, 25–60
cm), simple hairs especially in lower part and hispid (not
with only densely articulated glandular hairs or densely
glandular, sparsely bifurcate hairs below), hairs 2–2.5 mm
long (not ±1.5 mm), pedicel glabrous or sparsely bifurcate
and rarely trifid hairs (not only densely glandular hairs, or
bifurcate-stellate and glandular hairs), fruit glabrous (not
densely glandular, sparsely bifurcate hairs), valve narrower
than septum (not valve wider than septum).
Description: Perennial, rarely biennial herb. Roots
thickened taproot, (2.5–)4–16 mm diameter. Stem
±erect, 45–90(–110) cm tall (including flowering part),
generally purplish below or rarely completely, solitary or
2–4-branched flowering part, terete, smooth, 1.5–9 mm
in diameter at base; hispid especially in lower part, simple
hairs at bottom; bifurcate and simple hairs in middle part;
bifurcate, a few trifid, simple, and rarely glandular hairs
in upper part, long simple hairs 2–2.5 mm long. Leaves
crowded at bottom and middle part; basal leaves ±narrowly
lyrate, 5–10(–14) × 0.6–2(–3.5) cm (including 2.5–5 cm
petiole), remotely denticulate; cauline leaves decreasing
to flowering part, all with main midrib ±conspicuous;
lower cauline leaves bifurcate, glandular, a few simple and
3 branched hairs, simple hairs especially on main midrib
and branched hairs on margin, ±obtuse; middle cauline
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DURAN and ÇETİN / Turk J Bot
Figure 2. Stem, fruiting pedicel, and flowering pedicel features. Stem hairy: A- H. tosyaensis (A.Duran 5822), B- H.
ozcelikii (A.Duran 4636), C- H. bicuspidata (A.Duran 7443); fruiting pedicel (1- pedicel, 2- fruit): D- H. tosyaensis
(A.Duran 5822), E- H. ozcelikii (A.Duran 4636), F- H. bicuspidata (A.Duran 7443); flowering pedicel (1- pedicel, 2calyx): G- H. tosyaensis (A.Duran 5657), H- H. ozcelikii (A.Duran 5262), I- H. bicuspidata (Hamzaoğlu 2649).
Figure 3. Distribution map of Hesperis tosyaensis (■) and H. ozcelikii (●). A- Mediterranean
phytogeographical region, B- Euro-Siberian phytogeographical region, C- Iran-Turanian
phytogeographical region.
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Figure 4. Metaphase chromosomes of Hesperis tosyaensis
(A.Duran 5822) (2n = 14). Scale bar = 5 μm.
Coefficient
Figure 5. Dendrogram showing genetic relationships of some Hesperis species using ISSR markers.
leaves ±narrowly oblong, oblanceolate, lanceolate, or
lyrate, petiolate or sessile, attenuated at the base, serrate,
±acute, densely bifurcate, and sparse or a few simple and
3–4-branched hairs, simple hairs especially on the petiole
and main midrib, and branched hairs on the margin; upper
cauline leaves very narrowly linear-lanceolate, sessile
or rarely semiamplexicaule, slightly contracted at the
base, serrate, acute to acuminate. Inflorescence raceme,
rarely waxy panicle, branches ascending to erect, 10–45 ×
5–35 cm. All flowers ebracteate. Pedicels ascending, very
graceful, 12–22 mm long at anthesis, elongating to 28 mm
long at most, glabrous or sparsely bifurcate and rarely
trifid hairs. Sepals partly or entirely pinkish, inner sepal
oblong-oblanceolate, outer sepal oblanceolate-oblong,
deciduous, 5–8 veins, 7–8.5 × 1.4–2.2 mm, bifurcate, a
few trifid hairs and a few simple hairs only on tips, with
membranous margins, inner sepals strongly saccate.
Petals obovate or oblanceolate, 19–26 × 4.5–6 mm long,
purplish to violet, veins mostly conspicuous and purplish;
limb obovate, tapering gradually into the claw, 9–13 ×
4.5–6 mm, rounded, ±horizontal; claw 9–13 × 1.3–1.7
mm, clearly exerted sepal. Outer filaments not dilated at
base, 3.8–4.5 mm long, inner filaments dilated at base,
6.4–7.2 mm long, glabrous. Anthers all fertile, ±linear,
3–3.6 mm long, yellowish or greenish, basifixed. Stigma
with 2 obtuse, decurrent carpidial lobes. Ovary glabrous.
Fruiting pedicels not thickened towards the fruit, graceful,
0.4–0.5 mm in diameter. Siliquae (20–)40–70(–100) ×
1–1.5 mm, terete, dehiscent, torulose, straight or rarely
slightly curved, ascending to erect, glabrous, greenish
or partly purplish; valves slightly narrower than septum;
septum membranous or semimembranous in seed place,
remaining spongiose, with median veins inconspicuous.
Seeds ±dark brown, 2–2.6 × 1–1.3 mm, 1–24 in number.
Chromosome number: 2n = 14.
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(3867) are the same as H. tosyaensis. For this reason, the
specimen is called a paratype.
IUCN Red List category: The specimens were collected
in Kastamonu Province in Turkey, where the species seems
to be very rare and local, from an area of ca. 2.5 km2. The
population is not in good condition, with approximately
135 specimens. In the latest studies, this species was not
collected in Safranbolu and is only known from Tosya.
This species is evaluated as Critically Endangered [CR
B2ab (i-v)] (IUCN, 2010).
Distribution, habitat, and biogeography: Turkey is
the most complex country in the Middle East with regard
to geographic structure and landforms. It comprises
comparatively narrow and long, variously oriented
mountain chains separated by deep valleys and also highand medium-elevational plateaus. Central Anatolia is
predominantly a rolling plateau, the bulk of which ranges
from 900–1200 m in elevation. While more or less secluded
in the north, east, and south by higher mountain ranges,
it is open to wide valleys coming from the west (Zohary,
1973). Central Anatolia creates a barrier between the
Mediterranean and Black Sea regions. Hesperis ozcelikii,
which is a local endemic in southern Anatolia (Isparta
Province), is an eastern Mediterranean element, while
Hesperis tosyaensis appears to be endemic to the area of
Tosya (Kastamonu Province) and Safranbolu (Karabük
Province), northern Anatolia, and is a European-Siberian
element. Hesperis tosyaensis grows on roadsides, the edges
of stony piles, open places, and under open Quercus forest
at 900–1200 m (Figure 3).
Figure 6. Profiles of studied species on agarose gel, amplified
with ISSR primers. 1-, 2- Hesperis matronalis, 3- H. tosyaensis,
4- H. ozcelikii, 5- H. bicuspidata, 6- H. buschiana, 7- H. bottae,
8- H. pendula, 9- Matthiola anchoniifolia, 10- Hesperis hedgei, 11H. breviscapa.
Phenology: Hesperis tosyaensis has been found
flowering in May and June, and fruiting in June and July.
Paratypes: A5 Kastamonu: between Tosya-Sekiler
village, 7 km, 1000 m, 21.07.2001, under an open Quercus
forest, 40°55′62″N, 34°02′06″E, A.Duran 5822 & Menemen
(KNYA, ANK, HUB, GAZI); Paplagonia, Wilajet
Kastambuli (A4 Kastamonu): Tossia (Tosya) in fruticdis ad
Karkun, 24.05.1892, Sintenis 3867 (WU!, BM!, K!, ANK!);
Anatolia, Safranbolu, Wiedemann s.n. (K!, G!).
Specimen collected by Sintenis (3867) was identified as
H. armena by Prof C Haussknecht. After type specimens
of H. aspera were examined in Geneva Herbarium, it
was understood that the specimens collected by Sintenis
92
4. Discussion
Hesperis tosyaensis differs from H. ozcelikii by having its
stem erect, 45–90 cm (not ascending to erect, 25–60 cm),
hairs 2–2.5 mm long (not ±1.5 mm), inflorescence 10–45
× 5–35 cm (not 4–35 × 3–15 cm), pedicel very graceful
(not ±thick), glabrous or rarely sparse hairs (only densely
glandular hairs), ovary glabrous (not hairy), fruiting
pedicel 0.4–0.5 mm in diameter (not 1 mm in diameter),
fruit glabrous (not densely glandular, sparsely bifurcate
hairs), siliquae 1–1.5 mm wide (1.6–2.2 mm wide). More
comprehensive descriptions are given in Table 2.
Key to closely related Hesperis species
1. Perennial, rarely biennial herb, erect, 45–90 cm, hispid
hairs 2–2.5 mm long, inflorescence 10–45 × 5–35 cm;
pedicel glabrous or rarely sparse hairs, fruiting pedicel
0.4–0.5 mm diameter; fruit glabrous, valve narrower
than septum ............................................... H. tosyaensis
1. Biennial, ascending to erect, 25–60 cm, only densely
articulated glandular hairs, hairs ±1.5 mm long;
inflorescence 4–35 × 3–15 cm; pedicel densely glandular
hairs, fruiting pedicel ca. 1 mm in diameter; fruit densely
glandular, valve wider than septum ............. H. ozcelikii
DURAN and ÇETİN / Turk J Bot
Table 2. The comparison of diagnostic characters of Hesperis tosyaensis with H. ozcelikii.
Diagnostic characters
Hesperis tosyaensis
Hesperis ozcelikii
Hesperis bicuspidata
Life form
Perennial, rarely biennial
Biennial
Perennial
Stem height
Erect, 45–90 cm
Ascending to erect, 25–60 cm
Ascending to erect or erect, 20–50 cm
Stem
Generally purplish at bottom,
1.5–9 mm diameter at base
Rarely purplish at bottom, 3–6 cm diameter at base
Generally green, rarely purplish at bottom, 2–6
mm diameter at base
Stem hairs
Simple hair at bottom
Only densely articulated glandular hairs or densely
glandular and sparsely bifurcate hairs below
Densely short stellate-furcate and very rarely a
few glandular hairs in lower part
Hairs
2–2.5 mm long
±1.5 mm long
up to 1 mm
Leaves
Narrowly lyrate
Lyrate or sinuate dentate
Generally entire
Petiole
2.5–5 cm long
1–3 cm long
1.5–4 cm long
Inflorescence
Raceme, rarely waxy panicle,
branches 10-45 × 5–35 cm
Generally simple raceme, branches 4–35 × 3–15 cm
Generally raceme, sometimes branched at base,
5–20 × 5–20 cm
Pedicel
Very graceful, glabrous
±thick, only densely glandular hairs
±thick, short stellate-bifurcate and
rarely patently stalked glandular hairs
Pedicel length
12–22 mm long at anthesis,
elongating to 28 mm long at most
5–20 mm long at anthesis, elongating to 25 mm
long at most
4–12 mm long at anthesis, elongating to 16 mm
long at most
Anther
3–3.6 mm long
3.5–4 mm long
2.5–4 mm long
Ovary
Glabrous
Hairy
Glabrous or hairy
Fruiting pedicel
Graceful, 0.4–0.5 mm diameter
Slightly thickened, ±1 mm diameter
Slightly thickened, 0.6–1 mm diameter
Fruit wide
1–1.5 mm
1.6–2.2 mm
1.4–2.2 mm
Fruit
Glabrous
Densely glandular, sparsely bifurcate hairy
Valve
Narrower than septum
Wider than septum
Hesperis tosyaensis differs from H. bicuspidata by its
stem erect, 45–90 cm (not ascending to erect, 20–50 cm),
hairs 2–2.5 mm long (not up to 1 mm), leaves narrowly
lyrate (not entire), pedicel very graceful (not ±thick), 12–
22 mm long at anthesis (not 4–12 mm long at anthesis),
glabrous or rarely sparse hairs (not short stellate-bifurcate
and rarely patently stalked glandular hairs), fruiting pedicel
graceful, 0.4–0.5 mm in diameter (not thickened, 1 mm
in diameter), siliquae 1–1.5 mm wide (1.4–2.2 mm wide).
More comprehensive descriptions are given in Table 2.
Hesperis tosyaensis is closely related to Hesperis
buschiana, which is spread throughout northeastern
Anatolia. H. tosyaensis mainly differs from H. buschiana
in pedicel ascending, very graceful, 12–22 mm long
at anthesis (not ascending to erect, ±thick, 7–12 mm),
pedicel elongating to 28 mm long at most (not to 25 mm),
pedicel glabrous or sparsely bifurcate and rarely trifid
hairs (not bifurcate and trifid); petals 19–26 mm long (not
17–20 mm); fruiting pedicel not thickened towards the
fruit, graceful, 0.4–0.5 mm in diameter (not gradually and
Glabrous or asperous
Wider than septum
conspicuously thickened towards the fruit, 0.7–1.3 mm in
diameter).
Chromosome numbers of Hesperis taxa are reported as
n = 6, 7, 8, 12, 14, 16, 20, 24 and 2n = 12, 14, 16, 24, 28
(Manton, 1932; Khosravi and Maassoumi, 1998; Warwick
and Al-Shehbaz, 2006; Duran et al., 2008; Ünal et al., 2008).
The somatic chromosome number of Hesperis tosyaensis
was identified as 2n = 14 (Duran et al., 2008). Our results
are similar to those given in the literature (Figure 4).
The ISSR marker system is a PCR-based technique
that uses a single amplification primer composed of a
microsatellite motif to target a subset of simple sequence
repeats (SSRs) or microsatellites (Zietkiewicz et al.,
1994; Gürkök et al., 2013). ISSR-PCR is a simple, quick,
and efficient technique. It has high reproducibility. The
amplified products (ISSR markers) are usually 200–2000
bp long and amenable to detection by both agarose and
polyacrylamide gel electrophoresis (Reddy et al., 2002).
The technique is useful in areas of genetic diversity,
phylogenetic studies, gene tagging, genome mapping,
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DURAN and ÇETİN / Turk J Bot
and evolutionary biology in a wide range of crop species
(Reddy et al., 2002).
Twenty ISSR primers were used for PCR amplification.
Nine of them were chosen for ISSR analyses. The GC
percentages of selected primers were within the range of
33.3%–68.4%. A total of 104 bands were produced, of which
77 were polymorphic (74.03%). The total number of bands
ranged from 5 to 25. The lowest ISSR bands were produced
by primer M1. The highest ISSR bands were produced by
primer Cucurbita. Genetic similarity was estimated using
simple matching coefficients (Table 3). Genetic similarity
between studied species ranged from 0.5185 to 0.9814. The
highest similarity (0.9814) was observed between different
specimens of Hesperis matronalis. The genetic similarity of
H. ozcelikii and H. tosyaensis is 0.8518. Hesperis matronalis
has a high similarity (0.8518) to H. bicuspidata, H.
ozcelikii, and H. tosyaensis. The lowest similarity (0.4814)
was observed between Matthiola and Hesperis hedgei.
Six main clusters were observed in the dendrogram.
The first clade includes Matthiola. This species was used
as outgroup. The other clades include section Hesperis
taxa. The second clade includes Hesperis bottae Fourn.,
which is found in section Diaplictos (Dvorak) Dvorak.
Hesperis bottae is distinguished from other Hesperis taxa
by its sepals, petals, and stamens persistent in fruit. The
third clade incudes 2 subclades. The first subclade includes
Hesperis hedgei Davis, which is found in section Delicate
A.Duran. This taxon has the shortest calyx, corolla, and
fruit among Hesperis taxa. Section Delicate is related to
section Diaplictos. Section Delicate can be distinguished
from section Diaplictos by its fruit being glabrous, thinner,
and shorter, pedicel not thick and ascending. The second
subclade includes Hesperis breviscapa Boiss., which is
placed in section Cvelevia. All flowers of Hesperis breviscapa
are bracteate. Fruits are flattened siliquae that significantly
narrow towards the tip and open easily. The fourth clade
includes Hesperis pendula, which has pendulous fruit.
Hesperis pendula, which is widely distributed in all regions
of Turkey, is found in section Pachycarpos. The sixth clade
consists of the taxa of section Hesperis. Section Hesperis
is found spread across a wide geographic area, including
southern Europe and the Near East, from the mountainous
regions of southern Yakutia, western China, and across
Mongolia. The sections in northwestern Iran within Asia
Minor, the Balkan Peninsula, and the Caucasus are especially
rich (Tzvelev, 1959). The sixth clade contains 3 subclades,
the first of which is Hesperis buschiana (Figure 5). Hesperis
buschiana is endemic to Turkey and is known from only one
locality within the region of Artvin Province. The second
subclade includes Hesperis bicuspidata and H. matronalis.
Hesperis bicuspidata is a species endemic to Turkey and
is found in the Irano-Turanian phytogeographic region
in Turkey. Hesperis bicuspidata is the most widespread
species of Hesperis in Anatolia. There are wide variations
in plant height, leaf shape and size, calyx color and length,
corolla color, shape, size, venation and fruit hair, and shape
characters among the H. bicuspidata samples collected from
different localities. The third subclade includes Hesperis
tosyaensis and Hesperis ozcelikii. Results of molecular
analyses have confirmed that Hesperis tosyaensis is a distinct
species (see Table 3 and Figure 5). In conclusion, Hesperis
species studied in this manuscript show a correlation
between the morphological diagnostic characters and
molecular taxonomic classification in the dendrogram.
Table 3. Similarity matrix of studied species using ISSR markers.
Species
H. tosyaensis
H. tosyaensis
1
H. ozcelikii
0.8518
1
H. bicuspidata
0.8148
0.7777
1
H. buschiana
0.7592
0.7592
0.6851
1
H. matronalis
0.8333
0.8333
0.8333
0.7777
1
H. matronalis-1
0.8518
0.8518
0.8518
0.7962
0.9814
1
H. pendula
0.6851
0.6481
0.7222
0.6666
0.8148
0.7962
1
H. breviscapa
0.7777
0.7407
0.7037
0.6481
0.7222
0.7407
0.5740
1
H. hedgei
0.7037
0.6666
0.6666
0.6481
0.7222
0.7407
0.5740
0.7037
1
H. bottae
0.6111
0.6111
0.6481
0.5185
0.7037
0.7222
0.5555
0.5370
0.6111
1
Matthiola
0.5925
0.5185
0.6296
0.5370
0.6481
0.6666
0.6851
0.5555
0.4814
0.5370
94
H. ozcelikii
H. bicuspidata
H. buschiana
H. matronalis
H. matronalis-1 H. pendula
H. breviscapa
H. hedgei
H. bottae
Matthiola
1
DURAN and ÇETİN / Turk J Bot
As a result, comprehensive morphological studies
indicate that Hesperis aspera and H. bicuspidata cannot be
separated from each other as different species. According
to the priority rule (Greuter, 2000), H. aspera should be
treated as a synonym of H. bicuspidata. The specimens
of H. tosyaensis were examined and compared with
specimens of the related species H. ozcelikii in Turkey, and
Hesperis tosyaensis is described as a new species.
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Appendix.
Specimens examined (*also used for resources as
DNA samples): – Hesperis tosyaensis: A5 Kastamonu:
between Tosya-Sekiler village, 7 km, 1000 m, 21.07.2001,
under an open Quercus forest, 40°55′62″N, 34°02′06″E,
A.Duran 5822* & Menemen (KNYA, ANK, HUB, GAZI).
– Hesperis ozcelikii: Turkey. C3 Isparta: Sütçüler, 17th
km from Ayvalıpınar to Kesme, 1025 m, 37°36′53″N,
31°10′26″E, 11.06.1999, A.Duran 4636, H.Özçelik &
Sağıroğlu (KNYA); ibid., 04.08.1999, A.Duran 4973
(KNYA); ibid., 09.06.2000, A.Duran 5262* & H.Özçelik
(KNYA); Isparta: Sütçüler, between Ayvalıpınar-Kesme,
1300 m, 11.05.1997, H.Özçelik 7714 (GUL). – Hesperis
bicuspidata: Herbier de Mr. Benj. Delessert, ...., Aucher
109 (G) (type of Hesperis aspera); Tournefort, Nr. 16 (BM);
A4 Kastamonu: Ilgaz Mountain, Pinus sylvestris forest,
1500-2000 m, 30.06.1973, H.Peşmen & S.Erik s.n. (HUB);
A5 Yozgat: Aydıncık, Kazankaya town, Kazankaya canyon
pass, 650 m, 15.05.2000, stony places, shrubbery, A.Duran
5071 (KNYA); Kastambuli (Kastamonu): Tossia (Tosya),
Giaurdagh (Gavur Dağı), 10.06.1892, Sint. 3631 (WU,
P); A7 (Gümüsckhane) Gümüşhane: 28.05.1894, Sint.
5660 (K, WU, P); Gümüşhane: Köse, Kabaktepe village,
Kabaktepe, 2100 m, 16.07.2000, steppe, 40°15′94″N,
39°40′65″E, A.Duran 5551 (KNYA); Bayburt: between
Bayburt-Aşkale, Kop Mountain, 2300-2500 m, 31.06.1999,
stony steppe, 40°02’28″N, 40°30’22″E, A.Duran 4802 &
Akgül (KNYA); B4 Kırıkkale: Balışeyh, Mehmetbeyobası,
1350 m, 25.05.2001, Hamzaoglu 2649; B5 Kayseri: Felahiye,
Büyüktoraman village, Ak Dağ, Hamza Sultan tepe, 2100
m, 18.07.1999, steppe, A.Duran 4948 & Akgül (KNYA);
B6 Sivas: between Pınarbaşı-Gürün, 1695 m, 08.06.2007,
A.Duran 7443; B7 Erzinghan (Erzincan): Sipikordagh,
05.07.1889, Sint. 1201 (WU, K); B9 Bitlis/Van: 10 km
SE of Pelli, 8500 (2500 m), 8.06.1954, Davis 22597 &
Polunin (K); C2 Burdur: Dirmil-Korkuteli road, left of
Dirmil throat, 1650 m a.s.l., 09.06.1999, serpentine open
place, 36°58′08″N, 29°35′11″E, A.Duran 4628 (KNYA);
C3 Isparta: between Şarkikaraağaç-Yenişarbademli,
from Gedikli village to Çiçek Mountain, 1750–1850 m,
05.06.1999, A.Duran 4578* (KNYA); C6 Kahramanmaraş:
Tekne Da. between Göksun & Maraş, 1500 m, 04.05.1957,
rocky limestone slopes, Davis 27541 & Hedge (BM).
Synonyms of H. bicuspidata: – Hesperis stellata: Turkey.
A7 Gümüşhane: Prov. Pontus, distr. Gümüşhane, Kop
Dagh Pass, 2440 m a.s.l., 3 Aug 1960, rocky slopes, Furse
& Synge 865 (holotype: W, left-hand plant; isotype:
E!). – Hesperis tauricola: Iter cilicico-kurdicum, 5000′,
24.v.1859, Kotschy 180 (BM! E!). – Hesperis sintenisii:
Erzincan: Egin (Kemaliye), Bagir Başı, 3.5.1890, Sint.
2514 (holotype: JE; isotype: LD). – Hesperis kotschyana:
Turkey, in monte Tauro. Aestate, 1524 m, 24.05.1838
Kotschy 24 (holoype: K!). – Hesperis violacea: Turkey:
Mersina, 1843, Pinard s.n. (type: GOET!). – Hesperis
reuteriana: Turkey: prope İspir, in incultis, 6/1853,
Pavillon s.n. (type: FI photo! JE photo! BM!). – Hesperis
unguicularis: Turkey: environs d’Erzeroum, Calvert s.n.
(isotype: JE). – Cherianthus bicuspidatus: Turkey: Herbar
Gundelsheimer (type: B photo!). – Hesperis matronalis:
A6 Giresun: between Giresun-Şebinkarahisar, 44 km, 650
m, 15.07.2000, A.Duran 5539* (KNYA); Ardahan: Göle,
between Tellioğlu-Dörtkilise village, 2000 m, 27.06.1999,
A.Duran 4733* (KNYA); – Hesperis hedgei: C7 Şanlıurfa:
Siverek, northwest of Karacadağ village, 26.05.2000, 1100
m, 37°44′32″N, 39°38′91″E, A.Duran 5185* & Hamzaoglu
(KNYA). – Hesperis buschiana: A8 Artvin: Ardanuç,
Karlı village, 860 m, 01.06.2000, 41°05′62″N, 42°07′34″E,
A.Duran 5237* & Kandemir (KNYA). – Hesperis bottae:
B7 Elazığ: Harput, 21.06.1999, steppe, 38°42′80″N,
39°15′85″E, A.Duran 5182* & Hamzaoglu (KNYA). –
Hesperis breviscapa: B7 Erzincan: Üzümlü, Keşiş Dağı, 15
km, 2400-2500 m, 03.07.2014, 40°00′00″N, 35°09′37″E,
A.Duran 4825* & Akgül (KNYA). – Hesperis pendula: B5
Kayseri: Yahyalı, Aladağlar, 2600 m, 11.08.2001, A.Duran
5834 (KNYA). – Matthiola anchoniifolia: B6 Malatya:
between Darende-Gürün, 1260 m, 13.06.2007, A.Duran
7452* (KNYA).
1