Karyotypic study in some Iranian species and populations of Tulipa L. (Liliaceae)


Sheidai, M.; Shirin, Z.F.; Shadi, K.; Zehzad, B.

Caryologia 55(1): 81-89

2002


Karyotypic studies were performed on 12 populations of 7 Iranian tulip species (Tulipa biflora, T. sorgdiana, T. humilis, T. montana var. montana, T. montana var. chrysantha, T. systola, T. hoogiana) for the first time. The species/populations possessed 2n=2x=24 chromosome number but varied in details of karyotype including type of chromosomes, karyotype symmetry and number of SAT-chromosomes and differed significantly in total chromatin length as well as length of chromosome arms. A pair of heteromorph chromosomes occurred in T. humilis. The use of karyotypic data in taxonomy of the genus Tulipa is discussed.

CARYOLOGIA
Vol.
55,
no.
1:
81-89,
2002
Karyotypic
study
in
some
Iranian
species
and
populations
of
Tulipa
L.
(Liliaceae)
SHEIDAI
MASOUD*,
ZOGAGI-FAR
SHIItIN
,
KHANAFSHAR
SHADI
and
ZEHZAD
BAHRAM
Biology
Department,
Shahid
Beheshti
University,
Evin,
Tehran,
Iran.
Abstract
-
Karyotypic
studies
were
performed
on
12
species/
populations
of
Iran-
ian
tulips
for
the
first
time.
The
species/
populations
possessed
2n
=
2x
=
24
chro-
mosome
number
but
varied
in
details
of
karyotype
including
type
of
chromo-
somes,
karyotype
symmetry
and
number
of
SAT-chromosomes
and
differed
sig-
nificantly
in
total
chromatin
length
as
well
as
length
of
chromosome
arms.
A
pair
of
heteromorph
chromosomes
occurred
in
T
humilis.
The
use
of
karyotypic
da-
ta
in
taxonomy
of
the
genus
Tulipa
L.
is
discussed.
Key
words:
Cluster
analysis,
Heteromorph
chromosomes,
Iran,
Karyotype,
Tulips.
INTRODUCTION
Tulips
(Tulipa
L.)
are
among
important
plants
widely
used
as
ornamentals,
they
have
been
orig-
inated
in
Eastern
countries
and
via
Iran
and
Turkey
were
introduced
in
Europe
(MATIN
1998).
The
number
of
Tulip
species
occurring
in
Iran
varies
according
to
different
authors.
BoIssIER
(1882)
reported
seven
Tulipa
species
from
Iran
while
WENDELBO
(1977)
reported
12
species,
RECHINGER
(1990)
reported
19
and
Parsa
re-
ported
23
species
(MATIN
1998).
Tulipa
species
are
distributed
in
two
sub-gen-
era
namely
Eriostemones,
including
three
sec-
tions
and
Tulipa,
including
five
sections
(RAAms-
DONK
and
VARIES
1995).
Several
species
belong-
ing
to
these
sections
occur
in
Iran.
Due
to
eco-
nomic
importance
of
tulips,
they
have
been
sub-
jected
to
extensive
cytogenetical
studies
in
other
countries
(for
example
ATHANASIOU
1988;
JOHN-
SON
and
BRANDHAM
1977),
but
there
has
been
no
report
from
Iran.
*
Correspondin
g
author:
fax
++98
21
2403041;
e-mail:
msheidai@yahoo.com
The
present
paper
is
a
part
of
biosystematic
study
in
Tulipa
of
Iran
reporting
karyotypic
fea-
tures
in
some
of
the
species
for
the
first
time
try-
ing
to
present
the
chromosome
number
of
these
species
and
elucidate
the
karyotypic
changes
dur-
ing
the
species
diversification.
It
is
also
attempt-
ed
to
reveal
the
use
of
such
data
in
delimiting
the
species
studied.
MATERIALS
AND
METHOD
Karyotypic
studies
were
performed
in
12
popula-
tions
of
seven
Tulipa
species/
varieties
from
two
sub-
genera
of
1)
subg.
Eriostemones
including
Tulipa
bi-
flora
Pall
and
T
sogdiana
Bunge
from
the
sec.
Biflores
as
well
as
T
humilis
Herbert
from
the
sec.
Sazatiles;
2)
subg.
Tulipa
including
T
montana
Lindl.
var.
mon-
tana,T
montana
L.
var.
chrysantha
(Boiss.)
from
the
sec.
Clusianae,
T
systola
Stapf.
and
T
hoogiana
B.
Fedtsch.
from
the
sec.
Tulipanum.
Fresh
roots
obtained
from
collected
healthy
bulbs
were
used
for
cytological
preparations
using
2%
ace-
to-orcein
and
0.2
M
8-hydroxy
quinolin
(SHEIDAI
et
al.
2000b).
Chromosomes
were
identified
according
to
LEVAN
et
al.
(1964).
Karyotypes
were
compared
using
coefficient
of
variation
(CV;
AT
.
ERMA
1980),
total
form
percentage
(TF%;
HuzIwARA
1962),
Stebbins
,
two
is`e°
5'20
5'80
60
6 6
38
1
.
1
4
4
2
5
7
3
3
3
34
2
.
3
3
3
30'
1
.
26
1
ISLAMIC
REPUBLIC
OF
IRAN
S00
I
>
2000
82
SHEIDAI,
ZOGAGI-FAR,
KHANAFSHAR
and
ZEHZAD
way
system
of
karyotype
symmetry
(STEBBINS
1971)
and
Romero-Zarco
indices
(ROMERO
ZARCO
1986).
In
order
to
show
quantitative
differences
among
different
populations
of
a
single
species,
factorial
analysis
(ANOVA)
was
performed
on
the
size
of
the
chromosomes,
long
arms
as
well
as
short
arms,
using
the
chromosomes
and
the
species/
populations
as
the
two
factors
(SHEIDAI
et
al.
2000a).
In
order
to
compare
the
present
findings
and
those
reported
by
the
other
authors
on
similar
species
or
closely
related
ones
and
due
to
difference
in
cytological
preparation,
the
rela-
tive
values
of
karyotypic
features
were
used.
For
grouping
the
species/
populations
having
similar
karyotypic
features,
relative
data
were
stan-
dardized
(mean
=
0,
variance
=
1)
used
in
cluster
analysis
and
ordination
based
on
principal
compo-
nents
analysis
(PCA)
(SHEIDAI
et
al.
2000b).
Euclid-
ean/
squared
Euclidean
distance
was
used
as
a
mea-
sure
of
similarity
in
duster
analysis.
In
order
to
iden-
tify
the
most
variable
karyotypic
characters
among
the
species
studied,
factor
analysis
was
performed
(SHEIDAI
et
al.
1999).
Multivariate
statistical
analyses
used
SPSS
ver.
9.
(1988)
software.
RESULTS
AND
DISCUSSION
Geographical
distribution
of
Tulipa
species/
populations
studied
is
presented
in
Fig.
1
and
their
somatic
chromosome
number
as
well
as
karyotypic
datails
are
presented
in
Figs.
2
and
3
and
Tables
1
and
2.
All
the
species
and
popula-
Fig.
1—
Geographical
distribution
of
Tulipa
species
studied.
1=
Tulipa
montanum
var.
mon-
tanum,
2
=
T.
montanum
var.
crusantha,
3
=
T.
biflora,
4
=
T.
sogdiana,
5
=
T.
humilis,
6
=
T
hoogiana,
7
=
T
systola.
KARYOTYPES
OF
IRANIAN
TULIPS
83
tions
studied
possessed
2n
=
24
chromosome
number
supporting
the
earlier
reports
(UPCOT
and
LA
COUR
1936;
RAAMSDONK
and
VRIES
1992,
1995).
The
basic
chromosome
number
of
tulips
is
x
=
12,
therefore
the
species/
populations
studied
are
diploid.
However
triploid
(2n
=
36),
tetraploid
(2n
=
46)
as
well
as
pentaploid
(2n
=
60)
species
are
also
known
(RAAMSDONK
and
VRIES
1992,
1995).
The
occurrence
of
polyploidy
among
tulips
is
considered
as
a
mean
for
adaptation
to
envi-
ronmetal
variations
(BOTSCHANTZEVA
1982).
Although
the
species/
populations
studied
possessed
similar
chromosome
number,
they
dif-
fered
in
details
of
karyotypes
including
chromatin
length,
karyotypic
formulae
and
the
number
of
SAT-chromosomes
(Table
2).
Among
T
montana
populations,
the
lowest
value
of
total
haploid
chromatin
length
(133.8
pm)
occurred
in
Emamzadeh
Hashem
while
the
highest
value
(153.9
pm)
occurred
in
Abali
pop-
ulation.
Jajerood
population
possessed
the
high-
est
value
of
longest
chromosome
(20.6
pm)
while
Emamzadeh-Hashem
population
possessed
the
lowest
value
for
the
same
(15.5
pm).
The
highest
and
lowest
values
for
the
smallest
chromosome
occurred
in
Abali
(7.0
pm)
and
Jajerood
(6.0
pm)
populations
respectively.
Jajerood
population
possessed
the
highest
val-
ue
of
CV
(32.13)
showing
the
highest
variation
among
its
chromoromes
compared
to
the
other
T
montana
populations
while
Darabad
popula-
toin
possessed
the
lowest
value
(28.56).
These
populations
varied
in
their
karyotypic
formulae
too
(Table
2),
the
chromosomes
were
mostly
of
telocentric
and
sub-telocentric
and
1-2
sub-meta-
centric
chromosomes
except
in
Jajerood
popula-
tion
of
var.
chrysantha
which
possessed
only
st
and
t
chromosomes.
Variation
in
karyotypic
formulae
of
the
popula-
tions
studied
indicate
the
occurrence
of
structural
changes
in
the
chromosomes
which
is
supported
by
statistical
analyses
(following
paragraphs).
T
montana
populations
studied
occupied
4B
class
of
Stebbins
classification
which
is
among
ad-
vanced
classes
of
karyotype
symmetry
(more
asymmetrical).
Jajerood
population
due
to
its
Table
1-
Karyotypic
details
of
Tulipa
species/populations.
Species/
Population
Locality
Voucher
No.
2n
x
TL
Pm
L
pm
S
pm
L/S
pm
X
pm
Tulipa
var.
montana
montana
Golestan
99102
24
2x
147.8
17.5
6.7
2.6
12.3
T
montana
var.
montana
Emamzadeh-
Hashem
99105
24
2x
133.8
15.5
6.1
25
11.1
T
montana
var.
montana
Abali
99108
24
2x
153.9
20.0
7.0
2.8
12.8
T
montana
var.
crysantha
Darabad
99104
24
2x
147.6
17.5
6.4
2.7
12.3
T
montana
var.
crysantha
Jajerood
99109
24
2x
151.2
20.6
6.0
3.4
12.6
T
humilis
Touchal
99131
24
2x
162.1
20.3
9.4
2.1
13.5
T
humilis
Hamedan
99125
24
2x
198.6
22.6
11.4
1.9
16.5
T
In:flora
Tehran
2000-5
24
2x
217.1
24.4
13.4
1.8
18.1
T
sogdiana
Semnan
2000-7
24
2x
181.3
22.7
10.0
2.2
15.1
T
hoogiana
Golestan
99134
24
2x
184.7
20.8
10.3
2.0
15.4
T
systola
Bamu
2000-2
24
2x
198.4
22.3
11.7
1.9
16.9
T
systola
Darabad
99139
24
2x
198.4
22.3
11.7
1.9
16.5
Abbreviations:
x
=
Ploidy
level,
TL
=
Total
haploid
chromatin
len
g
th,
L
=
Lon
g
est
chromosome,
S
=
Shortest
chromosome,
L/S
=
Lon
g
est/
shortest
chromosome,
X
=
Mean
chromatin
length.
84
SHEIDAI,
ZOGAGI-FAR,
KHANAFSHAR
and
ZEHZAD
high
Al
value
(0.86)
and
low
TF%
(13.70)
pos-
sesses
the
most
symmetrical
karyotype
while
Golestan
population
with
a
lower
Al
value
(0.82)
and
higher
TF%
(17.70)
possesses
the
most
asymmetrical
karyotype.
Pearson
coefficient
of
correlation
determined
for
karyotypic
parametrs
revealed
a
high
value
(r
>
0.90)
for
total
chromatin
length
and
long
arm
of
the
chromosomes,
indicating
homogenity
of
the
karyotypes
and
inclusion
of
these
varieties
in
the
same
species.
However
a
lower
value
of
cor-
relation
for
short
arm
of
the
chromosomes
(r
<
0.60)
and
arm
ratio
(r
<
0.70)
indicates
changes
in
the
short
arms
as
well
as
arm
ratios
due
to
structural
changes
of
the
chromosomes
(SHEDAT
et
al.
2000b).
ANOVA
test
performed
on
the
size
of
chro-
mosomes,
long
arms
as
well
as
short
arms
among
T
montana
populations
showed
presence
of
sig-
nificant
difference
(p
<
0.05)
in
the
size
of
short
arms
only.
This
may
indicate
that
during
varieties
differentiation
or
population adaptation,
no
sig-
nificant
quantitative
change
has
occurred
in
total
and
the
long
arm
length
of
chromosomes,
but
a
significant
change
has
occurred
in
the
short
arms
length.
The
variations
observed
in
karyotypic
details
of
the
two
varieties
and
their
populations
as
well
as
lack
of
a
significant
difference
in
the
size
of
chromosomes
indicate
that
karyotypic
features
may
not
help
in
differentiating
the
varieties
from
each
other
and
morphological
characters
are
more
helpful.
Among
the
other
species
studied
Bamu
pop-
ulation
of
T
systola
possessed
the
highest
mean
total
chromatin
length
(202.33
mm)
and
the
low-
est
value
occurred
in
Tochal
population
of
T
hu-
milis.
The
highest
value
for
longest
chromosome
occurred
in
T
biflora
(24.40
mm)
while
the
low-
est
value
occurred
in
Touchal
population
of
T
hu-
milis
(20.37
mm).
The
highest
value
of
CV
occurred
in
T
sogdi-
ana
(24.76)
and
the
lowest
value
occurred
in
T
bi-
flora
(18.68).
Both
values
are
higher
than
those
observed
in
the
populations
of
T
montana
var.
montana
and
var.
crysantha.
Therefore
these
kary-
Table
2
-
Karyotypic
formulae
and
symmetry
(localities
as
in
Table
1).
Species
SAT
No.
SAT-
ch
TF%
CV
KF
ST-
class
Al
A2
T
montana
var.
montana
NO
NO
17.70
32.70
lsm
+
9st
+
2t
4B
0.82
0.33
T
montana
var.
montana
NO
NO
15.70
28.70
2sm
+
7st
+
3t
4B
0.83
0.29
T
montana
var.
montana
NO
NO
16.60
31.10
lsm
+
lOst
+
It
4B
0.83
0.31
T
montana
var.
crysantha
NO
NO
15.80
28.56
lsm
+
8st
+
3t
4B
0.83
0.29
T
montana
var.
crysantha
NO
NO
13.70
32.13
7st
+
5t
4B
0.86
0.32
T
humilis
NO
NO
23.10
24.20
lsm
+
llst
4B
0.68
0.20
T
humilis
3
1,
2,
3
19.80
20.30
2sm+
lOst
4A
0.72
0.20
T
biflora
2
2,
6
21.10
18.60
2sm+
lOst
4A
0.72
0.18
T
sogdiana
1
5
20.82
24.76
3sm+
9st
4B
0.72
0.24
T
hoogiana
2
11,
4
20.85
19.68
lsm+
llst
4B
0.75
0.19
T
systola
NO
NO
18.89
18.73
12st
4A
0.74
0.19
T
systola
NO
NO
19.98
19.29
12st
4A
0.76
0.18
Abbreviations:
SAT
No.
=
Number
of
SAT
chromosmes,
SAT-ch
=
Chromosomes
carrying
secondary
constriction,
TF%
=
Total
form
percentage,
CV
=
Coefficient
of
variation,
KF
=
Karyotypic
formulae,
ST-class
=
Stebbins
,
class,
NO
=
not
observed.
KARYOTYPES
OF
IRANIAN
TULIPS
85
otypes
show
more
variatoin
in
the
size
of
chro-
mosomes
compared
to
those
of
T
montana
vari-
eties.
Pearson
coefficient
of
correlation
determined
for
karyotypic
parameters
among
the
species
studied
revealed
a
high
value
(r
>0.90)
for
total
chromatin
length
and
long
arm
of
the
chromo-
somes,
indicating
homogenity
of
the
karyotypes
and
inclusion
of
the
species
in
the
same
genus.
However
a
lower
value
of
correlation
for
short
arm
of
the
chromosomes
(r
=
0.10
-
0.60)
and
arm
ratio
(r
=
0.40
-
0.70)
indicates
changes
in
the
short
arms
as
well
as
arm
ratios
due
to
structur-
al
changes
of
the
chromosomes.
ANOVA
test
performed
on
karyotypic
para-
meters
among
different
species
studied
revealed
b
d
e
f
O
C.
Altwolniwflipi
Fig.
2
Representative
somatic
cells
and
karyogram
in
Tulipa
species.
a
=
Tulipa
humilis
Hamedan,
b
=
T
hoogiana
Gorgan,
c
=
T
sogdiana
Semnan,
d
=
T
&flora
Tehran,
e
=
T
systola
Darabad,
f
=
karyogram
of
Tulipa
humilis
Hamedan
showing
a
heteromorph
bivalent.
Scale
=
10
pm.
I
25
MN
-
20
15
1
10
6
7
8
9
10
11
12
1
2
3
4
5
6
3
1
2
h
111
11
12
1
2
3
4
5
6
7
8
9
10
11
12
18
16
14
12
10
8
6
4
2
0
86
SHEIDAI,
ZOGAGI-FAR,
KHANAFSHAR
and
ZEHZAD
a
1111ii
4
5
6
7
8
9
10
11
12
miumi
11111111
2
3
4
5
6
7
8
9
10
11
12
1
2
3
25
20
15
10
5
0
c
d
i
No
m
Ili
_
10
11111111
5
1111
15
0
2
3
4
5
6
7
8
9
10
11
12
1
2
3
4
0
e
25
20
WM
7
8
9
10
11
12
f
9
MI
7
8
9
10
1111111
1
2
3
4
5
6
30
25
20
15
10
5
0
25
20
15
10
5
0
25
20
15
10
5
0
25
20
15
10
5
0
10
.
_
1111111
6
7
8
1
10
11
12
Fig.
3
Representative
ideograms
of
Tulipa
species.
Scale
=
pm.
a
=
Tulipa
biflora,
b
=
T
systola
Fars,
c
=
T
systola
Tehran,
d
=
T
humilis
Hamedan,
e
=
T
humilis
Touchal,
f
=
T
hoogiana,
g
=
T
sogdiana
h
=
T
montana
var.
montana
Emamzadeh
Hashem,
i
=
T
montana
var.
crysantha
Darabad.
20
15
10
5
0
1
2
3
4
5
KARYOTYPES
OF
IRANIAN
TULIPS
87
the
presence
of
a
significant
difference
(p
<
0.05)
in
total
chromosome
length,
long
arms
of
the
chromosomes
as
well
as
arm
ratios.
A
significant
difference
(p
<
0.05)
in
the
size
of
short
arms
of
the
chromosomes
occurred
only
between
T
bi-
flora
and
the
other
species
studied.
Therefore
karyotypic
changes
during
Tulipa
speciation
has
been
accompanied
with
significant
chromatin
changes
(either
loss
or
gain;
quantitative
change)
as
well
as
qualitative
change
as
is
revealed
in
dif-
ference
in
karyotypic
formulae
and
arm
ratios.
This
is
also
supported
by
differences
observed
in
the
karyotypic
formulae
and
SAT-
chromosomes
(number
of
SATs
and
the
chromosomes
carrying
secondary
constriction)
in
the
species
studied
(Table
2).
In
order
to
compare
the
present
findings
with
those
reported
from
the
other
species
of
the
subg.
Tulipa
from
former
USSR
(BOCTSCHANZEVA
1982),
the
relative values
of
karyotypic
features
were
used.
Stebbins
,
two
way
system
of
karyotype
classification
revealed
that
in
the
subg.
Tulipa,
T
systola
and
T
hoogiana
from
the
section
Tuli-
panum
occupy
4A
and
4A
and
4B
respectively,
T
montana,
T
wilsoniana
and
T
linifolia
from
the
section
Clusianae
occupy
4B
class
while
T
lehmanniana
(sec.
Kolpakovskianae),
T
miche-
liana
(sec.
Eichler)
and
T
scherenki
(sec.
Gesne-
rianae)
occupy
3B
class.
These
three
sections
are
also
considered
close
to
each
other
based
on
their
morphological
characetrs
(MATIN
1998).
In
the
subg.
Eriostemons
(sec.
Biflores),
T
bi-
flora
occupies
3A
class
while
T
sogdiana
occupies
4B
class.
Therefore
T
sogdiana
possesses
a
more
advanced
(asymmetrical)
karyotype
compared
to
T
biflora.
Populations
of
T
humilis
occupy
4A
and
4B
classes.
Therefore
in
general
Tulipa
species
(from
both
sub-genera)
possess
asym-
metrical
karyotype
as
they
occupy
mostly
4A
and
4B
classes
of
Stebbins
,
two
way
system
of
karyo-
type
classification.
From
the
present
results
it
seems
that
increase
in
the
karyotype
asymmetry
is
accompanied
with
the
loss
of
chromatin
material
as
the
species
with
more
asymmetric
karyotype
(4B
class)
possess
sig-
nificantly
shorter
total
chromatin
length
com-
pared
to
the
species
with
less
asymmetric
kary-
otype
(4A
class).
Therefore
chromatin
loss
may
be
one
of
the
possible
karyotypic
changes
during
the
species
diversification
in
the
genus
Tulipa.
Cluster
analysis
and
ordination
of
the
species/
populations
from
the
subg.
Tulipa
based
on
the
relative
karyotypic
data
produced
similar
results
(Figs.
4
and
5),
indicating
distinctness
of
the
species
studied
as
they
stand
in
separate
groups.
This
is
also
supported
by
ANOVA
test
performed
on
the
size
of
chromosomes
as
well
as
chromo-
some
arms,
indicating
a
significant
difference
among
the
species
studied.
An
interesting
result
is
the
separation
of
T
montana
from
T
linifolia
which
are
very
simi-
lar
in
morphological
characters
but
due
to
kary-
otypic
difference
stand
separate
from
each
other.
Therefore
karyotypic
features
may
help
in
differ-
entiating
the
two
species.
T
systola
and
T
hoogiana
from
the
section
Tulip
anum
are
placed
close
to
each
other
and
form
a
group,
while
T
micheliana
(sec.
Eichler)
is
joined
them
with
some
distance
followed
by
T
lehmanniana
(sec.
Kolpakovskianae).
The
same
relationship
is
revealed
from
morphological
characters
(MATIN
1998).
The
separation
of
T
scherenkii
from
the
other
species
is
also
similar
to
morphological
characters.
PCA
analysis
of
relative
karyotypic
data
re-
vealed
that
the
first
3
factors
comprise
about
68%
of
total
variance.
In
the
first
factor,
which
comprises
about
35%
of
total
variance,
the
rela-
tive
length
of
chromosomes
1,
3
and
5
possessed
the
highest
correlation
(>0.60)
and
are
the
most
variable
karyotypic
characters
of
the
first
factor.
This
factor
separates
T
wilsoniana
from
the
oth-
er
species.
In
the
second
factor
which
comprises
about
19%
of
total
variance,
the
relative
length
of
chromosomes
6,
7
and
9
possessed
the
highest
correlation
(>0.60).
This
factor
separates
the
populations
of
T
montana
and
T
scherenki
from
the
other
species
(Fig.
5).
It
is
intresting
to
mention
that
in
Hamedan
population
of
T
humilis
the
first
pair
of
chromo-
somes
was
heteromorph
and
differed
in
size
in
all
the
somatic
cells
studied
(Fig.
2).
Various
sources
have
been
suggested
for
the
occurrence
of
asy-
metrical
bivalents
including
breakdown
of
a
larg-
er
complex,
defeciencies
and
duplication
het-
erozygotes,
pericentric
and
paracentric
inversion
heterozygotes,
unequal
translocation
(SYBENGA
1992)
as
well
as
chromosome
fragmentation
(SHEIDAT
et
al.
2000a).
However
the
reason
for
the
occurrence
of
asymmeric/
heteromorph
chro-
mosomes
in
T
humilis
is
not
known
yet.
In
short
the
present
study
reveals
that:
1)
Tuli-
pa
species
of
Iran
possess
similar
chromosome
number
(2n
=
24)
and
such
data
is
not
useful
in
the
species
delimitation;
2)
the
species/
popula-
tions
vary
in
their
karyotypic
formulae
and
SAT-
Rescaled
Distance
Cluster
Combine
CASE
0
5
10
Label
Num
+
12
13
h
11
h
14
m
3
b
15
1
2
le
1
w
5
mc
6
rnm
8
mc
7
mc
9
mm
10
sch
4
3
2•
15
20
25
1•
MC
MC
mm
<
01
0
0-
-1
le
MC
mm
h
b
sch
-2.
-3
-20
-1.5
-1.0
-.5
PCA
2
1.5
88
SHEIDAI,
ZOGAGI-FAR,
KHANAFSHAR
and
ZEHZAD
Figs.
4
and
5
UPGMA
cluster
analysis
and
PCA
ordination
of
Tulipa
species.
Abbreviations:
b
=
T
biflora,
h
=
T
hoogiana,
1=
T.
linifolia,
le
=
T.
lehmania,
m
=
T.
micheliana,
mc
=
T.
montana
var.
crysantha,
mm
=
T.
montana
var.
mantana,
s
=
T.
sogdiana,
sch
=
T.
schrenkii.
KARYOTYPES
OF
IRANIAN
TULIPS
89
chromosomes
and
show
a
low
value
of
r
value
for
short
arm
of
the
chromosomes
and
arm
ratio
in-
dicating
the
occurrence
of
structural
changes
in
the
chromosomes;
3)
the
species
studied
differ
significantly
in
their
total
chromatin
length,
indi-
cating
the
occurrence
of
a
significant
gain/
loss
during
the
species
diversification;
4)
two
varieties
of
T
montana
var.
montana
and
var.
crysantha
do
not
differ
significantly
in
their
total
chromatin
length
and
show
inter-population
variation
in
karyotypic
details,
therefore
karyotypic
features
can
not
help
in
differentiating
the
two
varieties
from
each
other
and
morphological
characters
are
more
helpful.
However
these
two
varieties
are
distinct
in
their
karyotypic
features
compared
to
T
linifolia
which
are
otherwise
very
similar
in
morphological
characters.This
is
the
first
cytlog-
ical
report
in
Tulipa
species
and
populations
of
Iran.
REFERENCES
ATHANASIOU
K.,
1988
Some
cytogeographical
notes
on
the
genus
Tulipa
L.
(Liliaceae).
Botanika
Chronika,
8:
21-59.
BoissiER
E.,
1882
Flora
Orientalis.
Vol
V:
191-201.
Geneva
and
Basel.
BOTSCHANTZEVA
Z.P.,
1982
Tulips:
Taxonomy,
Morhology,
Cytology,
Phytogeography
and
Phys-
iology.
English
translated
edition
by
H.
Q,
Varekamp,
Rotterdom:
Balkema.
HuzIwARA
Y.,
1962
Karyotype
analysis
in
some
genera
of
Compositae.
VIII.
Further
studies
on
the
chromosomes
of
Aster.
Amer.
J.
Bot.,
49:
116-
119.
JOHNSON
M.A.T.
and
BRANDHAM
P.E.,
1977
New
chromosome
numbers
in
petaloid
monocotyledons
and
in
other
miscellaneous
angiosperms.
Kew
Bulletin,
52:
121-138.
LEVAN
A.,
FREDGA
K.
and
SANDBERG
A.A.,
1964
Nomenclature
for
centromeric
position
on
chro-
mosomes.
Hereditas,
52:
201-220.
MATIN
F,1998
Tulips
of
Iran.
Publication
of
Agri-
cultural
Research,
Education
and
Development.
Tehran,
Iran.
RAAMSDONK
L.W.D.
VAN
and
VARIES
T.
DE.,
1992
Biosystematic
studies
in
Tulipa
sec.
Erioste-
mones
(Liliaceae).
Plant
Syst.
Evol.,
179:
27-41.
RAAMSDONK
L.W.D.
VAN
and
VARIES
T.
DE.,
1995
Species
relationships
and
taxonomy
in
Tulipa
subg.
Tulipa
(Liliaceae).
Plant
Syst.
Evol.,
195:
13-44.
RECHINGER
K.H.,
1990.
Tulipa
L.
In:
K.H.
Rechinger
(Ed),
Flora
Iranica,
165:
76-103,
Graz.
ROMERO
ZARCO
C.,
1986.
A
new
method
for
esti-
mating
karyotype
asymmetry.
Taxon,
35:
526-
530.
SHEIDAI
M.,
NARENGI
Z.
and
KHATAMSAZ
M.,
1999
Karyotype
and
seed
protein
analysis
of
Lyci-
um(Solanaceae)
in
Iran.
Edinb.
J.
Bot.,
56:
253-
264.
SHEIDAI
M.,
ARMAN
M.,
MOHAMADI-SAEID
A.
and
ZEHZAD
B.,
2000a
Notes
on
cytology
and
seed
protein
characteristics
of
Aegilops
species
in
Iran.
The
Nucleus,
43:
118-128.
SHEIDAI
M.,
NASIRZADEH
A.
and
KHERADNAM
M.,
2000b
Karyotypic
study
of
Echinops
(Aster-
aceae)
in
Fars
Province,
Iran.
Bot.
J.
Linn.
Soc.,
134:
453
-
463.
STEBBINS
G.L.,
1971
Chromosomal
evolution
in
higher
plants.
Edwar
Arnold
Publisher.
SYBENGA
J.,
1992
Cytogenetics
in
Plant
Breeding.
Berlin,
Springer-Verlag.
UPCOT
M.
and
LA
CouR
L.,
1936
The
genetic
structure
of
Tulipa.
I.
A
chromosome
survey.
Jour-
nal
of
Genetic,
33:
237
-
254.
VERMA
B.N.,
1980
Karyotype
analysis
in
three
species
of
Rhizoclonium
Kutze.
Cytologia,
45:
433
-
440.
WENDELBO
P.,
1977
Tulips
and
Irisis
of
Iran
and
their
relatives.
Tehran,
Botanical
Institute
of
Iran.
Received
October
24,
2001;
accepted
December
3,
2001