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OT SİSTEMATİK BOTANİK DERGİSİ
THE HERB JOURNAL OF SYSTEMATIC BOTANY
Sahibi (Publisher)
Editör (Editor)
: Prof. Dr. Şinasi YILDIRIMLI
: Prof. Dr. Şinasi YILDIRIMLI
YAYıN KURULU (Editorial Board)
Herve M. BURDET
Andre CHARPIN
Prof. Dr. Avinoam DANIN
Prof. Dr. Musa DOGAN
Prof. Dr. Sadık ERİK
Prof. Dr. Yusuf GEMİCİ
Prof. Dr. Özden İNCEOGLU
Prof. Dr. Georgia KAMARI
Prof. Dr. Osman KETENOGLU
Dr. Jan KlRSCHNER
Dr. Gerald KREBS
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Dr. Elena MONASTERIO HUELIN
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ISSN 1300-2953
© OT Sistematik Botanik Dergisi, Ankara-TÜRKİYE
Ön Kapak (Front cover): Erysimum dirmilense Yıldırımlı & M. Dinç (Brassicaceae)
Foto: Ş. YıldırımlI
OT Sistematik Botanik Dergisi
15,2,125-136,
2008-ISSN 1300-2953
(Printed date: December 2008)
Morphological,
anatomical
and palynological
properties
of
endemic Onosma sintenisii Hausskn. & Bornm.
(Boraginaceae)
Serdal
MAKBUL*
& Zafer TURKMEN**
Osman
*
Faculty of Arts & Sciences,
Rize University,
Giresun
University,
***
****
Erzincan
Karadeniz
University,
Technical
Department
of
Department
of
28000, Giresun-Turkey
Faculty of Education,
University,
Biology,
&
531 00, Rize-Turkey
Faculty of Arts & Sciences,
Biology,
DEMİR***
BEY AZOGLU****
Biology,
**
Ali KA
&
24030,
Erzincan-Turkey
Faculty of Arts & Sciences,
Department
of
61 080 Trabzon-Turkey
Abstract
In this
study,
(Boraginaceae)
palynologically.
endemic
was
Onosma
investigated
bases.
consists
of 3-5
layered
phloern.
The leaves are equifacial
or anisocytic.
The plant
morphologically,
anatomically
is 10-35 cm tall and erect.
collenchyrna
and
and
cambium
and amphistomatous.
Pollen type of species
axis and shape of the pollen is subprolata
Key Words:
is clear
Stomata
The stem
below
the
cells are
The stomata index is 21.5 for the upper epidermis
and 12.8 for the lower epidermis.
one ofpolar
& Bornm.
Hausskn.
Stern and leaves are covered with densely hairs. lt has setae
with tuberculate
anomocytic
sintenisli
Onosma sintenisii, morphology,
is syncolporatae
(P/E:
anatomy,
ı .2).
palynology
in
OT 15,2,
126
125- 136,2008
Endemik Onosma sintenisii Hausskn. & Bornm. (Boraginaceae)
türünün
morfolojik,
anatomik
ve palinolojik
özellikleri
Özet
Bu
çalışmada,
endemik
iBoraginaceaeı
morfolojik,
Gövde
ve yapraklar
sahiptir.
yoğun
Onosma
anatomik
tüylerle
Bitki 10-35 cm boyunda
mevcuttur
ve
kambiyum
ekvifasiyal
ve
amfısutomatiktir.
anizositiktir.
Sutoma
i2.8'dir.
kaplıdır.
indeksi
olarak
Tabanıarı
ve diktir. Gövdede
altında
Sutoma
Bomm.
incelenmiştir.
şişkin setalı tüylere
3-5 sıralı kollenkima
belirgindir.
hücreleri
Yapraklar
anomositik
için 2 ı.5, alt epidermis
üst epidermis
Onosma
sintenisii,
veya
için ise
(P/E: 1.2).
ve polen şekli subpurolatdır
Sözcükleri:
&
Hausskn.
ve palinolojik
fuloyemin
Polen tipi sirıkolporat
Bulduru
sintenisii
morfoloji,
anatomi,
palinoloji
Introduction
Onosma L. (Boraginaceae)
comprises
about
Turanian
regions
endenıic
150 species,
is the largest genera of the family (I). The genus
distributed
(2-4). In Turkey
mainly
in the Mediterranean
there are 95 species
(I). Sonıe new species have been deseribed
and Irano-
of Onosma, 48 of which are
and several new records
made
(5,6).
Riedl
(7) and
Boissier
(8) performed
so me organizations
Onosma using hairs characters.
Also
basal
in the Onosma species
cells
accounts
of the family
nıorphological
for Iran (II).
the inıportant
studied
of trichonıes
have been evaluated
characters
Erdtman
differ
Khatoon
and several
et aL. indicated
traits in the family.
in terrns of anatomical,
ecological
species
and palinological
mainly
as new records
that po Ilen characters
Some
of
taxonomic
(I O) employing
have been recognized
(12) and Diez (13) reported
taxononıic
that the nature
(9). Recently
by Khatamsaz
species
on the genus
are one of
of the Onosma were
properties
by Akçin
s. MAKBUL,
Z. TURKMEN, A. KANDEMİR, O. BEY AZOGLU
and Engin (14,15)
caryological
and Binzet
studies
and Orcan (16). Teppner
on the genus. Phenolic
species were studied
by Sanıhan
substituents
Anatomical
Watson
studies
and Dallwitzs
not specifically
on the genus
(23) explained
for the genus.
Metcalfe
the anatomical
emzik (Kemah,
Erzincan)
Onosma species
Hausskn.
been previously
of
is an endemic
reported
o.
(Kemaliye,
the morphological,
yalancı
(setae)
o.
and dyes (9, 26).
(VU) category
species
sintenisii
(27) that has
(28, 29). The anatomic
Therefore,
palynological
havacıva,
(25). Some
in Turkey
plant and a haplotrichous
in the Vulnerable
that hairs
but
within Onosma species.
Erzincan)
fo lk medicines
(22) and
of the family
(24) noted
otu (Eskişehir),
sinıenisii have not been studied.
paper is to investigate
of
and emcek
and Chalk
properties
and Chalk
as emzik
are used as herbs,
ex Bornm.
properties
are named
Onosma
of some endemic
types vary in Boraginaceae and it shows great variation
Onosma species
so me
et aı' (21).
are limited.
Metcalfe
(17, 18) performed
studies on the genus Onosma
et aı' (19). Chemical
carried out by Özgen et aL. (20) and Kundakovic
127
the purpose
and anatomical
of this
properties
o. sinıenisii.
Materials and Methods
Plant samples
Turkey.
Specimens
herbarium
Faculty
materials.
appearance
these
results
manually.
entellan
in Erzincan
specimens
were
were
compared
with
dried
University
characters
were
from B7 square
used
according
pollen
the flora
in
to standard
Herbarium
were measured
for
(Erzincan)
of Education
on the herbarium
studies.
of Turkey
Observed
(27).
General
of the plant was drawn.
For anatomical
sections
and stored
studies
Some morphological
AIso
morphological
sinıenisii were collected
for morphological
techniques
(EHEF).
o.
of
of upper
analysis,
and
These sections
(30). Photographs
and the stomata
lower
cross-sections
leaf surface
were stained
of stern and leaves,
were
used.
with safranin
All sections
as deseribed
were
taken
and fast green and fixed with
of them were taken with an Olympus
index was calculated
and superficial
by Meidner
BXSl
microscope
and Mansfield
(31).
OT 15,2, 125-136,2008
128
The po Ilen slides were prepared according to the Wodehouse technique (32).
PolIen grains were dissected from herbarium specimens on a clean microscope
slides and added 2-3 drops of 96 % ethanol to melt the resin and oil. GlycerinegelIatine added basic fuchsine was placed on pollen and allowed to melt and mixed
by a clean pin to get scattered
polIen grains
photographed on camera Olympus BX5
ı from
then the po Ilen shape were
permanent sı ide. The measurements
are based 30 readings from this species. Pollerı diameter,
polar axis (P) and
equatorial diameter (E), porus length and width, exine and intine thickness were
measured
under
light microscope.
Also ornarnentation
of pollerı grains was
determined. The terminology used is in accordarıce with Erdtman (12), Wodehouse
(32), Faegri and Iversen (33).
Results
Morphological Properties (Figure 1)
Perennial, caespitose.
setose and sparsely
Stern 2 or numerous,
pubescent,
adpressed
setose above,
narrowly
linear-lanceolate
10-35 cm, simple, adpressed
base clothed with remains
sparsely
setose beneath:
long petiolate,
of leaves. Leaves
basal 30-150 x 3- ıO mm,
cauline and bracts linear-Ianceolate,
Inflorescerıce terminal and lateral subcapitate
of 1-4 cymes. Pedicel to 3 mm.
Calyx 13- i 6 mm in flower to 18 mm in fruit, lobes broadly lanceolate,
acute,
widening in fruit and soon glabrescent. Corolla yellow, 20-22 mm, campanulate,
hairy. Filaments inserted near base of corolla, slightly longer than included anthers.
Nutlets c. 3.5 mm, oblong-ovoid, acute, brown, smooth or slightly rugose. Fl. 5-7.
Examined specimens: Turkey. B7 Erzincan: İliç, around Hasanova viIIage,
gypsum steppe, 1023 m, 12.vı.2004, A. Kandemir 6064; İliç, Kuruçay, gypsum
steppe, 1220 m, ll.vı.2005,
A. Kandemir 6950.
S. MAKBUL, Z. TURKMEN, A. KANDEMİR, O. BEY AZOGLU
129
~)
\'
c
L..J
b
Lı mm
e
Figure I. a. General
of Onosma sintenisli Hausskn.
appearance
Kandemir 6950) b. Corolla with stamens,
c. nutlet, d. indumentum,
& Bomm. (A.
e. setae on leaf
Anatomical Properties
Stem (Figure
Epidermis
rectangular
hairs
underneath
was observed
or compressed
abundantly
collenchyma
2)
in transactions.
cells. Theyare
on the epidennis.
tissues
consisting
the epidermis.
located under the collenchyma.
The
4-6
covered
Theyare
of 3-5
it composed
layers
layered
of a single
with a thin cuticle.
unicellular
There
or multicellular.
of rectangular
compressed
layer of
cells
parenchyma
are
are
The
located
tissue
is
OT 15,2, 125-136,2008
130
Figure 2. Onosma sintenisli Hausskn. & Bornm. Transeetion of the stern, e:
epidermis, ep: cortex parenchyma, cl: collenchyma, phl: phloem, xl: xylem, h: hair,
cb: cambium, p: pith parenchyma
Diameter of parenchyma cells are 25-30 x 55-60 J.l and thickness of cortex is
approximately
300 u. Phloem ring (50 u) is composed
Cambium is distinguishable.
trachea cells are 10-28
jı.
of 4-5 layers of cells.
Xylem makes up 20-25 % of stem and diameter of
The pith consists of large and cylindrical parenchymatic
cells and makes up 50-60 % of stem (Tabı e 1).
Leaf (Figures 3-8)
The transverse seetion of the lamina, midrib and surface preparations of both
epidermises revealed the following elements. Midrib is triangle-shaped and has J-2
layered collenchyma located below the both epiderınal cells. Upper epidermal cells
are larger than the lower once. Hairs are present only on upper epidermis surfaces
in cross
sections.
Solitary
arc-shaped
vascular
bundles
are surrounded
by
parenchymatous and orbicular shaped bundle sheath. Diameter of these cells is 7080 u. Cambium is seen clearly below the phloem.
The leaves are equifacial. Palisade parenchyma cells are 3 layered (170-180
u) on the upper surface and 2 layered (90- ı00 u) on the lower surface. Spongy
parenchyma
cells with large intercellular
cavities are 3-4 layered (90 u). Both
epidermal cells have unduIate walls. Stomata cells are anisocytic and anomocytic.
S. MAKBUL, Z. TURKMEN, A. KANDEMİR, O. BEYAZOGLU
13]
The stomata occur on the both surfaces at the same level with neighboring cells.
Stomata index is 21.5 for the upper epidermis and 12.8 for the lower epidermis.
There are 489 stomata cells and 2285 epidermis cells on the upper surface and 408
stomata cells and 2775 epidermis cells on the lower surface (1 mm").
There are abundantly
tuberculate
setae and setules
Tuberculate setae have cystoliths in the bases and theyare
on the epidermis.
more common on the
upper epidermis. Setae occur only on upper epidermis surface. Both epidermis
surfaces covered witb setules (Table
ı, 2).
l~f1ın
pp
e
Figure 3-4. Onosma sintenisli Hausskn. & Bornm. Transections of the leaf. upe:
-U
upper epidermis, cl: collenchyma, le: lower epidermis, p: parenchyma,
pbl:
plıloem, xl: xylem, cb: cambium, vb: vascular bundle, lı: bair, pp: palisade
parenchyma, sp: spongy parencbyma
5
20
ın
6
Figure 5-6. Surface sections oftbe lower epidemıa. Stl: setule
100/:!..!!,l
OT 15,2, 125-136,2008
132
7
2~m
Figure 7-8. Surfaee seetions of the upper epiderma. st: setae, stl: setule
Table i. Anatomical measurements of Onosma sintenisii Hausskn. & Bornm.
Breadth (LL)
Min
Max
15
20
35
25
110
3
25
30
17
20
Epidermis cells
Collenchyma cells
Parenchyma cells
Diameter of trachea
Diameter of pith cells
Cuticle
Upper epidermis cells
Lower epidermis cells
Parenchyrna cells
Diameter of trachea
Stem
Leaf
25
25
40
30
130
5
30
35
20
25
Lerıs th (LL)
Min
Max
20
35
50
35
40
60
-
-
-
-
-
-
50
55
50
20
60
65
65
25
Table 2. Features of stomata and epiderma cells on the upper and lower epiderma of
Onosma sintenisii Hausskn. & Bornm.
Upper surface ofleaf
Number
Number
Stomata
Stomata
Stomata
of stomata (I mm")
of epiderma cells(l mm")
length (LL)
width (LL)
index
489
2285
40
28
21.5
Lower surface of leaf
408
2775
37
25
12.8
Palinological Properties (Figures 9-10)
Pollerı grains of Onosma sintenisii are monad, small, eireular, syneolporatae,
teetum; euteetate. Pollen diameter (Amb) is 13-14.53 ı-ı, polar axis (P) is 20-23.09
S. MAKBUL, Z. TURKMEN, A. KANDEMiR, O. BEYAZOOLU
u, equatorial
axis (E) is 16-20 fl and P/E rate is 1.15-1.25
is subprolata
in equatorial
between
1.27-1.44
4.25-4.45
u. Sexine
Orrıamentation
view and İs circular
fl and porus
is granulated
u. Nexine
u. Shape of pollen grains
in polar view. Porus length (plg) İs
width (plt) is 3.48-3.78
is 0.61-0.7
133
is 0.75-0.85
or slightly reticulates
u. The exine thickness
u. Intine
is 0.78-1.1
Figure 9-10. Pollen morphology
Equatorial
10
of Onosma sinıenisii Hausskn.
view;
& Bornm. 9.
i O. Polar view
Table 3. Pollen eharaeters of Onosma sintenisii Hausskn. & Bornm.
Amb
P
E
P/E
Plg
PIt
Exine
Sexine
Nexine
Intine
Pollen Shape
Omamentation
Po Ilen Type
Minimum
13
20
16
1.15
u.
(Table 3).
9
Average
13.69
21.65
17.88
1.2
Maximum
14.53
23.09
20
1.25
4.45
4.25
3.48
1.27
4.35
3.63
l.35
0.61
0.75
0.78
0.65
0.8
0.94
Suberolata
Granulate/slizhtlv retieulate
svncolnoratae
3.78
1.44
0.7
0.85
1.1
İs
OT 15,2,125-136,2008
134
Discussion
In this study, Onosma sintenisii collected from Hasanova-Erzincan
the type
locality of the plant were investigated in terms of anatomical, morphological
palynological
Turanian
properties.
element.
Examined
species is an endemic
The morphological
properties
and
taxon and Iranian-
determined
in our studyare
generally similar to those in flora of Turkey, but there are some differences among
the two studies. Stern is 10-28 cm, basal leaves glabrous beneath, basal 3S-1 00 mm
long and 3-6.S mm wide, shortly petiolate, inflorescence terminal, 1-2 cymes, calyx
lS-16 mm, and corolla glabrous, nutlets c. 3.S mm in the flora of Turkey (27). In
our study, stems 10-3S cm, basalleaves
beneath sparsely setose, basa120-70 x 2.S-
10 mm, cauline leaves 2S-4S x 2-3 mm, cauline leaves long petiolate, inflorescence
terminal and lateral, 1-4 cymes,
bracts were 1-2.1 x 0.1S-0.6 cm, the calyx 8-1S
mm in flowers and lS-23 mm fruit, the corolla 18-2S mm, hairy, and nutlets 2.S2.9 x 1.8-2 mm.
There are tuberculate setae on the epidermis of the stern and leaf. Setae have
12-1S tuberculate
tuberculates
hairs and cystoliths
is S-6 in Onosma
in
isauricum
o.
sintenisii.
The number of these
Boiss. & Heldr.,
8- 12 in Onosma
stenolobum Hausskn. ex H. Riedl and 7 (6) in Onosma bornmuelleri Hausskn. (14,
IS).
Metcalfe and Chalk (22) and Watson and Dallwitz (23) explained that the
xylem was composed
obviously.
of sclerenchymatic
The same results
Collenchyma
cells and trachea
were seen in
o.
and cambium
o.
roussaei,
giganteum
is
(6).
is 3-S layered in our study but some Onosma species have 1-3
layered collenchyma tissue (13). Palisade parenchyma generally is 3-layered on the
upper surface and 2-layered on the lower surface in our study.
o.
2(-3)-layered palisade cell on the upper surface and monolayered
bracteosum has
on the lower
surface (IS).
The leaf is equifacial. Palisade parenchyma cells are 3 layered on the upper
surface and 2 layered on the lower surface. However,
o.
bracteosum
has 2-3
layered palisade cells on the upper surface and single layered on the lower surface
S. MAKBUL,
(14).
The
Z. TURKMEN,
stomata
were
A. KANDEMiR,
anisocytic
and
O. BEY AZOGLU
anomocytic.
pointed out that there were both anomocytic
Metcalfe
and anisocytic
The same results were seen in so me other Onosma species
Onosına pollens are very easily distinguished
by its heteropolar,
species
shape,
shows
colpi
similar
pollen
syncolporatae
tricolporate
considerable
length,
tectal
grains
in
of
o.
surface
grains,
nıostly
Qureshi
(22)
in this family.
(14, 15).
fronı other types in the fanıily
in their general
(34).
stomata
Chalk
syncolpate.
pollen
and Qaiser
Onosma. The pollen
Generally,
the
characters
such
(35)
reported
grains
also
as
are cornpletely
o. sintenisii.
by Johnston
that
Echium, l.obosıemon,
(36). These findings
bracıeosum are subprolate
shape of
o. sintenisii
Alkanna.
Onosma have 3-
are in line with our study. Pollen grains
and theyare
sphaeroidea
in
o.
muıabile. Pollen
is subprolate.
With this study,
o.
variation
in the genus
lt was deterrnined
aperturate
pollen
and
135
nıorphological,
sintenisli was defined.
As conclu
pollen features were deternıined
anatonıical
and palinological
ion hair characters
as inıportant
infornıation
and anatonıical
properties of
traits with
for the genus.
References
i.
2.
3.
4.
5.
6.
7.
8.
9.
LO.
ii.
12.
YI LDl RI M Ll Ş., The chorology of the Turkish species of Boraginaceae family, Ot Sistematik
Botııııik Dergisi, 7. 2, 257-272, 2000.
TUTIN L HEYWOOD W. H., (eds.), Flora of Europaea, Cambridge Univ. Press., 3: 89-94,
1972.
ZOHARY M., Boraginaceae, in "Flura of Palestina", Jerusalern, 3: 70-72, 1978.
RIEDL H., Boragiııaceae in K.H. Rechinger (ed.) "Flora Iranica", 48,192, Graz, 1967.
RIEDL H., BİNZET R., ORC AN N.. A new speeies of On05111a (Boraginaccae-Lithospernıeaeı
Irorn souıhcm Turkey, Edinburgh Journal of Botany, 61, 127-130,2005.
BİNZET R., ORCAN N., Onosına sintenisli Hausskn. ex. Bornm. ve Onosına bulboırichuııı
DC iBoraginaceaeı
türlerinin palinolojik özelliklerinin belirlenmesi, 18. Ulusal Biyoloji
Kongresi, Aydın, 2006.
RIEDL H., Onosma, in Davis, P.II. (ed.) "Flora of Turkey and the East Aegean Islands", 6:
326-376, Edinburgh University Press, Edinburgh 1979.
BOISSI ER E., Flora Orientalis, Gerıevae, Basiliae, 3: 1810-1885, 1879.
KHATOO
S., MEHROTRA S., BAJPAI V.K., MEHROTRA B.. , Ulıramorphology ofsome
Boraginaccous ıaxa used as Raıanjoı, Feddes Repertorium,
i OS, 1-2,61-7, 1994.
KHATAMSAZ M., Onosına straııssii as a new combination and some new plant records in
Iran, i ran Journal of Borany, 5, 2, 75-82, 1992.
CIIAIIREMA
A .. ATTAR F" Ane\\' species of the genus Onosına L. from W. Iran, Iran
Journal of Borany, 7. i. 51-57, 1996.
ERDTMAN G., Pollen ı\lol'phology and Plant Taxonomy-Angiospcnn
(in Introduction
to
Patynologv). \ ol. i. Almquist and Wiksell. Stockholm. 1952.
136
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
OT IS, 2, 125-136, 2008
DIEZ M.J., Contribucion
al atlas polinico de Andalucia Occidental,
i. Boraginaeeae,
Lagascalia, 13, 147-171, 1984.
AKÇiN Ö.E., E Gl A., Endemik Onosnıa bornmulleri Hausskn. 'nın morfolojisi, anatomisi
ve ekolojisi üzerine bir araştırma, Ekoloji, 13,51, 13-19,2004.
AKÇiN Ö.E., E GİN A., The rnorphological, arıatomical and ecological properties of endemic
Onosına bracteosuın Hausskrı. & Bornrn. (Boraginaeeae) species, Turk. J. Bot., 29, 317-325,
2005.
BiNZET R., ORCAN
., Morphological and palynological studies on Onosına roussaei DC.
and Onosma giganıeum Lam. (Boraginaeeae), Ot Sistematik Botanik Dergisi, i O, 57-76,
2003.
TEPPNER H., Karyosystematik von Onosma sıellulaıum, o. pygmaeum und O. lepıanıhum
(Boraginaeeae), Bot. Jahrb. Syst., 102 (1-4), 297-306,1981.
TEPPNER H., Onosına kaheirei spec. nova und o. eree/um (Boraginaeeae) aus Griechenland,
Phyton (Austria), 28, I, i 15-131, 1988.
SARUI-IAN N., KADIOGLU A., TÜRKMEN
Z., BEYAZOGLU
O, Doğu Karadeniz
Bölgesi'nde yayıfış gösteren bazı Onosma (Boraginaceae) türlerindeki
fenolik madde ve şeker
içeriklerinin araştırılması, XVii. Ulusal Biyoloji Kongresi, Adana, Türkiye, 2004.
ÖZGEN U., COŞKUN M., KAZAZ c., SEÇEN H., Naphthoquinones
from the roots of
Onosına argenıatum Hub.-Mor. (Boraginaeeae), Turk. J. Chem., 28, 451-454, 2004.
KUNDAKOVIC
T., FOKIALAKIS
N., DOBRIC S., PRATSINIS
H., KLETSAS D.,
KOVACEVIC N., CHINOU 1., Evaluation of the anti-inflammaıory and cytotoxic activities of
naphthazarine derivatives from Onosma leptantha, Phytomedicine,
1-5,2004.
METCALFE C.R., CHALK L., Anatomy of Dicotyledons
II, Oxford University Press,
London, 1979.
WATSON L., DALLWITZS, M.J., The families of angiosperrn: automated descriptions, with
interactive identifıcation and information retrieval, Aust. Syst. BoL., 4,68 i-695, i99 I.
METCALFE C.R., CHALK L., Anatomy of Dicotyledons I, Oxford University Press, London.
1950.
AKÇiN Ö.E., Engin A., Onosına isauricum ve o. sıenolobuın türlerinin karşılaştırmalı
anatomisi. Ot Sistematik Botanik Dergisi, 8, 2, 75-95, 200 I.
ÖZTÜRK M., ÖZÇELiK H., Doğu Anadolu'nun
Faydalı Bitkileri, Ankara, SÜSKA V 1991.
DA ViS P.H. (ed.), Flo1'3 of Turkeyand
the East Aegean Islands, vol. 6, Edinburgh
University Press, Edinburgh, 1978.
EKiM T., KOYUNCU M., VURAL M., DUMA
H., AYTAÇ Z., ADIGÜZEL N., Türkiye
Bitkileri Kırmızı Kitabı, Eğrelti ve Tohumlu Bitkiler (Red Data Book of Turkish Plants,
Pteridophyta and Sperınaıoptıytaı,
Ankara, Türkiye Tabiatı Koruma Derneği Yayınları, No:
18,2000.
i UCN, Red List Categories, version 3.1., prepared by the IUCN Species Survival Cornrnission.
Gland, Switzerland, and Cambridge, UK: IUCN, 200 i.
VARDAR Y., Botanikte Preperasyon Tekniği, Ege Üniversitesi, Fen Fakültesi Yayınları, No:
2, ızmir, 1987.
MEINDER H., MANSFIELD T.A., Physiology ofStomata, Graw-Hill, New York, 1968.
WODEHOUSE R. P., Pollen grains, New York, Hafner, 1935.
FAEGRI K., lVERSEN J., Text Book of Pollen Analysis, 2"d ed., Munksgaard, Coperıhagen,
1964.
PERVEEN A., SHAHEE
U., QAISER M., Pollen flora of Pakistan-IV, Boraginaceae, Pak. J.
Bot., 27, 2, 327-360,1995.
QURESHI U.S., QAISER M., Palynological study of Onosına (Boraginaceae) from Pakistan,
Pak. J. Bot., 19,99- i05, 1987.
JOHNSTO
R.F., Population
Structure
in Salt Marsh
Song Sparrows:
Part
i.
Environment and Annual Cycle, i: 58, pp. 24-44. 1954.
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