Species relationships and taxonomy in Tulipa subg. Tulipa (Liliaceae)


van Raamsdonk, L.W.D.; de Vries, T.

Plant Systematics and Evolution 195(1/2): 13-44

1995


Principal component and canonical variate analyses were used to analyse the variation in 34 morphological characters measured in 98 accessions of Tulipa subgenus Tulipa, formerly known as section Leiostemones. In addition, 43 accessions were analysed cytogenetically by means of C-banding methods. Low levels of heterochromatin (<2%) was found in sections Kolpakowskianae and Eichleres, while high levels (>20%) were present in some species of sections Tulipanum and Tulipa. Based on the encountered variation and on geographical distribution data, 30 species in 5 sections were recognized, of which the synonymy is stated. Seven new series in 2 different sections are designated.

—Plant
.
Systematics
and
Evolution
©
Springer-Verlag
1995
Printed
in
Austria
Pl.
Syst.
Evol.
195:
13-44
(1995)
Species
relationships
and
taxonomy
in
Tulipa
subg.
Tulipa
(Liliaceae)
L.
W.
D.
VAN
RAAMSDONK
and
T.
DE
VRIES
Received
April
15,
1994;
in
revised
version
October
28,
1994
Key
words:
Liliaceae,
Tulipa,
subg.
Tulipa,
sects.
Clusianae,
Kolpakowskianae,
Tulipa-
num,
Eichleres,
Tulipa.
Typification,
morphology,
numerical
taxonomy,
chromosome
banding,
biogeography,
nomenclature.
Abstract:
Principal
component
and
canonical
variate
analyses
were
used
to
analyse
the
variation
of
34
morphological
characters
measured
in
98
accessions
of
Tulipa
subg.
Tuli-
pa,
formerly
known
as
sect.
Leiostemones.
In
addition
43
accessions
were
analysed
cyto-
genetically
by
means
of
C-banding
methods.
Based
on
the
encountered
variation
and
on
geographical
distribution
data
30
species
in
five
sections
were
recognized,
of
which
the
synonymy
is
stated.
Seven
new
series
in
two
different
sections
have
been
designated.
The
first
intergeneric
subdivision
of
the
genus
Tulipa
L.
was
made
by
DE
REBOUL
(1847).
This
division
was
incomplete
and
consisted
primarily
of
species
known
locally
in
southern
Europe.
Later
REGEL
(1873)
and
BAKER
(1874)
produced
more
elaborated
subdivisions.
The
two
main
groups
recognized
by
REGEL
(1873)
were
named
Eriostemones
and
Leiostemones
(BoisslER
1882),
and
they
appeared
to
be
clearly
separated
from
each
other.
This
main
subdivision
is
still
in
use.
Some
sec-
tions
designated
by
BAKER
(1874)
were
placed
in
these
main
groups
(BoissiER
1882).
Sect.
Tulipa
(Leiostemones
Borss.)
is
mainly
found
in
the
primary
gene
center
in
Central
Asia
and
comprises about
40
species.
The
systematic
position
of
several
species
is
not
clear.
The
cultivated
tulip,
T
gesneriana
L.,
is
part
of
a
group
of
closely
related
species
in
this
section
with
crossability
relationships
(VAN
EIJK
&
al.
1991).
The
species
of
the
other
section,
Eriostemones,
migrated
from
the
primary
as
well
as
from
the
secondary
gene
center
(Caucasus)
westwards
into
western
Europe.
The
approximately
20
species
of
this
section
are
separated
into
three
well
divided
subsections
(VAN
RAAMSDONK
&
DE
VRIES
1992).
These
results
are
mainly
based
on
morphological,
crossability,
chromosome
and
chemical
studies
(NIEuw-
HOF
&
al.
1990,
VAN
EIJK
&
al.
1991,
VAN
RAAMSDONK
&
DE
VRIES
1992).
This
paper
deals
with
a
study
aimed
at
systematic
enumeration
of
species
of
sect.
Tulipa
based
on
morphological
and
cytogenetical
characteristics
and
on
data
of
geographical
distribution.
14
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
Table
1.
List
of
Tulipa
accessions.
Accession
number,
chromosome
number,
source
and
origin
are
mentioned.
Ploidy
levels
are
indicated
for
each
species.
*
Additional
accessions
used
for
completing
geographic
and/or
cytogenetic
data
but
not
included
in
the
morphological
analyses.
All
taxa
are
indicated
on
the
species
level
with
the
nomenclaturally
correct
name
Sect.
Clusianae
T.
aitchisonii
A.
D.
HALL,
2X
66336
source
unknown
71337
2n
=
24
C.
GREY-WILSON
&
R.
F.
HEWER
GWH
972:
S
of
Kabul,
Afghanistan
72103
2n
=
24
GIBBONS
S
0725
Al
73119*
2n
=
24
originally
from
Elburz
Mts,
Iran
76171
2n
=
24
clone
of
71336:
C.
GREY-WILSON
&
T.
F.
HEWER
GWH
935:
W
of
Bamian,
Afghanistan
76172
2n
=
24
clone
of
71337
77279
clone
of 72129:
A.
ALA,
Iran:
Tasham
valley,
Iran
79152
2n
=
24
clone
of 75307:
Jammu,
Kashmir
T.
clusiana
DC.
(including
f.
chrysantha);
4X,
5
X
65152
VAN
EEDEN,
Noordwijk,
Netherlands;
cv.
Cynthia
68590
2n
=
48
Thoolen,
Overveen,
Netherlands
70639
W.
KOOIMAN,
Enkhuizen,
Netherlands
71328
2n
=
60
C.
GREY-WILSON
&
T.
F.
HEWER
GWH
638:
N
of
Ghazni,
Afghanistan
75303
2n
=
48
Jammu,
Kashmir
T.
montana
LINDLEY,
2X
70607
2n
=
24
W.
KOOIMAN,
Enkhuizen,
Netherlands
74428
2n
=
24
originally
Kopeth
Dagh,
Turkmenistan
78394
2n
=
24
clone
of
68065:
Hort.
Bot.
Tashkent,
Uzbekistan
T.
linifolia
REGEL,
2
X
67261
2n
=
24
C.
G.
VAN
TUBERGEN,
Haarlem,
Netherlands
70596
2n
=
24
W.
KOOIMAN,
Enkhuizen,
Netherlands
73460
source
unknown
T.
maximowiczii
REGEL,
2X
65470
2n
=
24
J.
G.
VAN
EEDEN,
Noordwijk,
Netherlands
T.
batalini
REGEL,
2X
64181
2n
=
24
C.
G.
VAN
TUBERGEN,
Haarlem,
Netherlands
70597
2n
=
24
W.
KOOIMAN,
Enkhuizen,
Netherlands
70611
2n
=
24
W.
KOOIMAN,
Enkhuizen,
Netherlands
71328A
2n
=
24
C.
GREY-WILSON
&
T.
F.
HEWER
GWH
638:
No
of
Ghazni,
Afghanistan
73116
2n
=
24
originally
from
Elburz
Mts,
Iran
77282
clone
of
73119:
originally
from
Elburz
Mountains,
Iran
Sect.
Kolpakowskianae
T.
altaica
PALL.
ex
SPRENGEL
82105*
Hort.
Bot.
Moscow,
Russia:
E.
Kazakhstan
T.
kolpakowskiana
REGEL,
2X
(4X)
68596
2n
=
24,
48
Thoolen,
Overveen,
Netherlands
74228*
2n
=
24
W.
KOOIMAN,
Enkhuizen,
Netherlands
T.
iliensis
REGEL,
2X
74226
2n
=
24
W.
KOOIMAN,
Enkhuizen,
Netherlands
T.
anisophylla
VVEDENSKY,
2X
73458*
(received
as
T.
korshinskyi)
76333*
2n
=
24
Hort.
Bot.
Dushanbe,
Tadzhikistan:
W
Pamir
Alai
T.
ferganica
VVEDENSKY,
2X
74247
2n
=
24
Hort.
Bot.
Tashkent,
Uzbekistan
75116
VAN
TUBERGEN,
Haarlem,
Netherlands
T.
lehmanniana
MERCKLIN,
2X
77161
M.
H.
HooG,
Haarlem,
Netherlands
Tulipa
subg.
Tulipa
15
Table
1
(continued)
T
zenaidae
VVEDENSKY,
2X
79161
2n
=
24
clone
of
77299:
Z.
M.
SILINA,
Petersburg,
Russia
T.
tetraphylla
REGEL,
4X
65302
2n
=
48
Hort.
Bot.
Alma-Ata,
Kazakhstan
67368
2n
=
48
Hort.
Bot.
Petersburg,
Russia
T.
behmiana
REGEL,
4
X
84280
2n
=
48
Hort.
Bot.
Salaspils,
Lithuania
T.
ostrowskiana
REGEL,
4X
68055
2n
=
48
Hort.
Bot.
Moscow,
Russia
70600*
2n
=
48
W.
KOOIMAN,
Enkhuizen,
Netherlands
74102
2n
=
48
Hort.
Bot.
Tashkent,
Uzbekistan
78120*
2n
=
48
W.
KOOIMAN,
Enkhuizen,
Netherlands
Sect.
Tulipanum
T.
oculus-solis
ST.
AMANS,
2X
73109*
2n
=
24
T.
F.
HEWER:
Zagros,
Iran
77310
2n
=
24
W.
KOOIMAN,
Enkhuizen,
Netherlands
84259*
2n
=
24
P.
VISSER,
St.
Pancras,
Netherlands;
originally
from
Turkey
84260*
2n
=
24
P.
VISSER,
St.
Pancras,
Netherlands;
originally
from
Turkey
86221
L.
SCHOORL,
Lisse,
Netherlands
T
systola
STAPF,
2X
79154
2n
=
24
clone
of
77105:
Golestan
forest,
Iran
T.
stapfii
TURRILL,
2X
71341
Mrs
A.
ALA,
Iran:
Golestan
forest
near
Tangehgol,
Iran
76175*
clone
of
71341
77270
2n
=
24
clone
of
71340:
Mrs
A.
ALA,
Iran:
Golestan
forest,
Iran
81282
hybrid
between
T.
stapfii
(76175)
and
T.
praecox
(77101)
T
ulophylla
WENDELBO,
2X
71339*
Mrs
A.
ALA,
Iran:
Dashted
Nuzir,
Iran
76173
clone
of
71339
77269
clone
of
71339
T.
kuschkensis
B.
FEDTSCHENKO,
2
X
76167
2n
=
24
clone
of
71320:
C.
GREY-WILSON
and
T.
F.
HEWER
GWH
494;
N
of
Merat,
Afghanistan
T.
julia
C.
KOCH,
2X
68075
Hort.
Bot.
Tashkent,
Uzbekistan
72119
2n
=
24
Hort.
Bot.
Yerevan,
Armenia:
Monastery
Gegard
T.
aleppensis
BOISSIER
ex
REGEL;
2
X
,
3X
84257
P.
VISSER,
St.
Pancras,
Netherlands
84268
2n
=
36
W.
KOOIMAN,
Enkhuizen,
Netherlands
T.
praecox
TENORE,
3
X
67741*
2n
=
36
Station
Alpine,
Lautaret,
France:
Le
Chet,
Maurienne
valley,
Savoy,
France
77101
2n
=
36
H.
Q.
VAREKAMP,
Wageningen,
Netherlands:
Korc,u1a,
Croatia
83209
M.
H.
HOOG,
Haarlem,
Netherlands
Sect.
Eichleres
T.
lanata
REGEL;
2
X
,
3
X
65369
2n
=
36
C.
G.
VAN
TUBERGEN,
Haarlem,
Netherlands
76332*
2n
=
24
Hort.
Bot.
Dushanbe;
originally
Darvas,
Turkmenistan
T.
ingens
HOOG,
2X
70599
W.
KOOIMAN,
Enkhuizen,
Netherlands
74429
2n
=
24
M.
H.
HOOG,
Haarlem,
Netherlands:
Kopeth
Dagh,
Turkmenistan
T.
eichleri
REGEL,
2X
67234
2n
=
24
W.
KOOIMAN,
Enkhuizen,
Netherlands
67235
2n
=
24
C.
G.
VAN
TUBERGEN,
Haarlem,
Netherlands;
cv.
Excelsa
16
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
Table
1
(continued)
T.
micheliana
HooG,
2X
78356
2n
=
24
clone
of
71315:
C.
GREY-WILSON
&
T.
F.
HEWER
GWH
391:
NE
Iran
T.
tubergeniana
HOOG,
2X
65391
2n
=
24
C.
G.
VAN
TUBERGEN,
Haarlem,
Netherlands
T.
hoogiana
FEDTSCHENKO;
2X,
3X
69169
C.
G.
VAN
TUBERGEN,
Haarlem,
Netherlands
81107*
2n
=
24
Hort.
Bot.
Askhabad:
Kopeth
Dagh,
Turkmenistan
82116*
2n
=
36
P.
VISSER,
St.
Pancras,
Netherlands
T
fosteriana
HooG
ex
W.
IRVING,
2X
71324A
2n
=
24
C.
GREY-WILSON
&
T.
F.
HEWER
GWH
493: N
of
Merat,
Afghanistan
82331
VELDHUIZEN,
Hillegom,
Netherlands;
cv.
Madame
Lefeber
T.
greigii
REGEL,
2X
78122
2n
=
24
M.
H.
HOOG,
Haarlem,
Netherlands:
Samarkand
79163
2n
=
24
clone
of
77293:
Z.
M.
SILINA,
Petersburg,
Russia
T
mogoltavica
M.
POPOV
&
VVEDENSKY
78389
Z.
M.
SILINA,
Petersburg,
Russia
T.
alberti
REGEL,
2X
68061
2n
=
24
Hort.
Bot.
Tashkent,
Uzbekistan.
T.
vvedenskyi
BOCHANTZEVA,
2X
65249
Hort.
Bot.
Petersburg,
Russia
68087*
2n
=
24
Hort.
Bot.
Tashkent,
Uzbekistan
80290
2n
=
24
clone
of
76124:
ELLIOT,
Ashford,
Kent,
UK;
from
Tashkent,
Uzbekistan
(received
as
T
butkovii
81120
L.
SCHOORL,
Netherlands
86115
W.
KLETZING,
Germany:
Choros,
elev.
2300
meter,
Pamir
Alai
T.
butkovii
BOCHANTZEVA,
2
X
76124
2n
=
24
Hort.
Bot.
Tashkent,
Uzbekistan
T
sosnovskyi
AKHVERDOV
&
MIRZOJEVA,
2X
67356
Hort.
Bot.
Yerevan,
Armenia
72120
2n
=
24
Hort.
Bot.
Yerevan,
Armenia:
Megri,
Armenia
T.
praestans
HOOG,
2X
70626
2n
=
24
M.
H.
HooG,
Haarlem,
Netherlands
71331
C.
GREY-WILSON
&
T.
F.
HEWER
GWH
709:
Salang
pass,
N.
of
Kabul,
Afghanistan
76115
B.
ANDREASEN,
Denmark;
from
Hort.
Bot.
Oxford,
UK
77266
clone
of
71331
80285
clone
of
71331
T.
kaufmanniana
REGEL,
2X
65301
2n
=
24
Hort.
Bot.
Alma
Ata,
Kazakhstan
65252
Hort.
Bot.
Petersburg,
Russia
68014
Hort.
Bot.
Moscow,
Russia
T.
amblyophylla
(POST)
FEINBRUN,
2X
74111*
2n
=
24
Hort.
Bot.
Jerusalem:
Negev
desert,
Israel
T.
tschimganica
BOCHANTZEVA,
2X
68085
2n
=
24
W.
KOOIMAN,
Enkhuizen,
Netherlands
68086
Hort.
Bot.
Tashkent,
Uzbekistan
76123*
2n
=
24
ELLIOT,
Ashford,
Kent,
UK
T.
anadroma
BOCHANTZEVA,
2X
78395
2n
=
24
clone
of
68068:
Hort.
Bot.
Tashkent,
Uzbekistan
T.
dubia
VVEDENSKY,
2X
78386
2n
=
24
clone
of
77160:
M.
H.
HooG,
Haarlem,
Netherlands
80298*
2n
=
24
clone
of
77344:
W.
Tien
Shan
Mts
T.
subpraestans
VVEDENSKY,
2X
68083
2n
=
24
Hort.
Bot.
Tashkent,
Uzbekistan
Tulipa
subg.
Tulipa
17
Table
1
(continued)
Sect.
Tulipa
T
armena
Boiss.
,
2X
74223
2n
=
24
C.
G.
VAN
TUBERGEN,
Haarlem,
Netherlands
71
galatica
FREYN,
3
X
84266
2n
=
36
W.
KOOIMAN,
Enkhuizen,
Netherlands
T.
hungarica
BORBAS,
2X
66066
2n
=
24
Hort.
Bot.
Sofia,
Bulgaria
76181
clone
of
74197:
originally
from
Roman,
Romania
T.
rhodopea
VELENOVKSY,
2X
66078
Hort.
Bot.
Sofia,
Bulgaria
75102
Hort.
Bot.
Budapest,
Hungary
75108*
2n
=
24
Hort.
Bot.
Sofia,
Bulgaria
T.
didieri
JORDAN,
2
X
68007
2n
=
24
Hort.
Bot.
Gottingen,
Germany
T.
planifolia
JORDAN,
2
X
67737 2n
=
24
Station
Alpine,
Lautaret,
France:
Villars-Clement,
Maurienne
valley,
Savoy,
France
T.
marjolettii
PERR.
&
SONG.,
2X
66040
Hort.
Bot.
Mtinchen,
Germany
73120
2n
=
24
C.
G.
VAN
TUBERGEN,
Haarlem,
Netherlands
T.
grengiolensis
THOMMEN,
2
X
67738
2n
=
24
Station
Alpine,
Lautaret,
France:
Grengiols,
Wallis,
Switzerland
T.
suaveolens
ROTH,
2X
65038
2n
=
24
Hort.
Bot.
Rostow/Don,
Russia
T.
gesneriana
L.
78392
source
unknown
82334
Hort.
Bot.
Balcik,
Bulgaria
Material
and
methods
Material.
Sect.
Tulipa
is
raised
to
the
level
of
subgenus
for
reasons
as
discussed
in
the
second
paragraph
of
the
discussion.
This
indication
will
be
used
throughout
the
paper
in
order
to
reach
uniform
terminology.
120
accessions
of
54
taxa
of
subg.
Tulipa
used
are
summarized
in
Table
1,
together
with
their
ploidy
level
and
their
origin.
Vouchers
were
deposited
in
WAHO.
Each
accession
consists
either
of
a
population
propagated
by
seed
or
a
clone,
in
most
cases
propagated
from
a
population
which
was
collected
originally
in
the
wild.
However,
not
all
data
on
locality
are
available.
Data
on
geographic
distribution
of
each
species
were
taken
from
HALL
(1940),
HooG
(1973),
and
BOCHANTZEVA
(1982).
Names
of
species
considered
to
be
closely
related
with
the
species
used
(HALL
1940,
BOCHANTZEVA
1982)
are
taken
into
account
in
order
to
discuss
probable
conspecificity.
Multivariate
analyses.
Of
the
total
of
120
accessions,
98
were
used
for
multivariate
morphological
analysis.
The
characters,
all
measured
on
the
flowering
date,
are:
flower-
ing
date,
plant
length,
stem
diameter,
stem
pubescent
(y/n),
number
of
leaves,
width
of
second
lowest
leaf,
leaf
undulation
(none,
weak,
strong),
position
of
second
lowest
leaf
(prostrate,
intermediate,
erect),
leaf
pubescent
(y/n),
leaf
margin
ciliate
(y/n),
number
of
flowers,
length
of
outer
tepal,
width
of
outer
tepal,
width
of
inner
tepal,
blotch
length
at
center
of
tepal,
blotch
length
at
margin
of
tepal,
width
of
blotch
border,
occurrence
of
sec-
ondary
blotch
(y/n;
1989
dataset
only),
streak
width
at
dorsal
side
of
tepal,
base
of
tepals
bearded
(yin),
outer
side
of
tepal
pubescent
(y/n),
tip
of
outer
tepal
pubescent
(y/n),
tip
of
inner
tepal
pubescent
(y/n),
anther
length,
filament
length,
filament
colour
contrasting
with
flower
colour
(y/n),
stigma
hairy
(y/n).
18
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
The
characters
measured
after
lifting
of
the
bulbs
are:
hairy
coronet
at
base
of
bulb
(y/n),
hairs
at
lower
part
of
bulb
tunic
(none,
sparse,
strong,
woolly),
hairs
at
middle
part
of
bulb
tunic
(none,
sparse,
strong,
woolly),
hairs
at
upper
part
of
bulb
tunic
(none,
sparse,
strong,
woolly),
bulb
tufted
at
top
(y/n),
colour
of
bulb
tunic
(cream,
light-,
mediated-,
dark-brown),
number
of
offset
bulbs.
The
classes
of
discrete
characters
are
mentioned
in
brackets.
These
characters
have
been
measured
on
an
ordinal
scale.
All
other
characters
were
measured
on
a
continuous
scale.
Four
plants
of
each
accession
(two
repeats
with
two
plants
each)
were
measured
in
1988
[subsects.
Clusianae,
Kolpakowskianae
and
Tulipanum
(Oculus-solis)]
and
in
1989
[subsects.
Eichleres,
including
Tulipanum
sensu
HALL
and
Spiranthera,
and
Tulipa
(Ges-
nerianae)].
The
correlation
coefficients
between
all
characters
were
calculated.
The
char-
acter
pairs
width
of
outer
and
inner
tepal,
blotch
length
at
center
and
margin
of
tepal,
tepal
outer
side
pubescent
and
bearded
base
of
tepals,
and
hairs
at
middle
and
upper
part
of
bulb
tunic
each
were
related
empirically.
It
can
be
assumed
that
these
characters
are
under
the
same
genetic
control.
These
relations
were
approved
numerically
by
high
correlation
coefficients
(R
>
0.8).
One
character
of
each
pair
was
removed
from
the
final
dataset
in
order
to
limit
a
too
high
level
of
redundancy
and
to
avoid
an
effective
character
weight
of
2
for
an
attribute.
The
character
of
each
pair
which
showed
the
lowest
correlation
coeffi-
cient
with
the
remaining
characters
was
removed.
Therefore
the
characters
width
of
inner
tepal,
blotch
length
at
margin
of
tepal,
bearded
base
of
tepals,
and
hairs
at
middle
part
of
bulb
tunic
have
been
left
out.
The raw
data
matrices
containing
30
or
29
characters
in
the
1988
and
1989
dataset,
respectively,
were
standardized
prior
to
performing
the
final
analyses.
Multivariate
analy-
ses
were
carried
out
using
the
IRIS
program
package
(VAN
RAAMSDONK
1988).
Principal
component
analysis
was
calculated
based
on
a
correlation
matrix
with
equal
weight
allo-
cated
to
each
character.
Principal
components
are
newly
calculated,
compound
axes
rep-
resenting
the
maximum
amount
of
linear
variation
in
each
direction
of
the
multidimen-
sional
space.
The
relation
between
the
characters
and
the
principal
components
is
expressed
by
means
of
factor
loads,
the
objects
are
projected
onto
the
newly
calculated
axes
by
means
of
component
scores.
The
species
as
assigned
in
literature
are
indicated
by
different
signs
in
the
figures
of
the
principal
component
plots.
The
regions
of
newly
combined
species
as
designated
in
the
current
paper
are
indicated
by
straight
polygons.
In
some
other
cases
regions
of
single
species
are
encircled
when
necessary.
Cytotypes
are
indicated
by
broken
lines
(Fig.
3).
The
position
of
a
hybrid
population
is
indicated
by
a
dotted
line
(Fig.
5).
Fifteen
populations
representing
the
variability
in
all
five
subsections
were
studied
in
both
years,
which
makes
it
possible
to
transform
the
1988
data
into
the
1989
ranges
by
means
of
linear
regression.
Most
discrete
characters
remained
untransformed.
Five
char-
acters
showed
a
too
large
year
effect
to
be
included
in
the
combined
dataset.
Using
the
combined
dataset
with
24
characters
Canonical
variate
analysis
was
carried
out
based
on
within
and
between
groups
covariance
matrices.
Canonical
variates
are
new
compound
axes
maximizing
the
distinguishing
ability
of
character
combinations
between
groups.
Chromosome
banding.
The
method
for
C-banding
of
chromosomes
was
briefly
as
follows:
root
tips
were
sampled
in
colchicine,
fixed
in
Carnoy
and
stored
at
—20
°C.
A
single
root
tip
was
macerated
in
45%
acetic
acid
at
room
temperature,
squashed
in
acetic
acid
and
air
dried
for
several
days.
C-banding
was
achieved
by
incubation
in
Ba(OH)
2
(3
min,
60
°C)
and
in
a
2x
SSC
solution,
cooling
from
60
°C
to
room
temperature
for
several
hours.
After
soaking
the
slides
were
stained
in
a
10%
Gun
solution
for
10
min.
More
details
on
the
C-banding
method
were
published
by
VAN
RAAMSDONK
(1984).
The
sensibility
of
the
dif-
ferent
accessions
showed
fluctuation,
so
minor
modifications
of
the
general
method
per
accession
had
to
be
used.
Tulipa
subg.
Tulipa
19
Table
2.
Percentage
of
explained
variation,
and
the
factor
loads
and
the
ranking
in
paren-
theses
of
the
15
main
determining
characters
for
the
first
four
principal
components
in
the
1988
dataset
with
sects.
Clusianae,
Kolpakowskianae,
and
Tulipanum.
*
Characters
with
high
factor
loads
exclusively
in
1988
First
PC
28.07%
Second
PC
10.53%
Third
PC
7.90%
Fourth
PC
6.53%
Stem
diameter
0.743
(3)
-0.201
(20)
0.106
(22)
-0.212
(11)
Stem
pubescence
0.255
(25)
0.460
(6)
-0.290
(12)
-0.543
(2)
Leaf
position
2nd
leaf*
0.437
(18)
0.326
(10)
0.584
(2)
0.247
(9)
Leaf
undulation
-0.147
(27)
-0.359
(9)
-0.471
(3)
0.148
(16)
Leaf
pubescence
0.319
(24)
0.185
(21)
0.027
(28)
-0.593
(1)
Width
of
second
leaf
0.878
(1)
0.125
(26)
-0.081
(24)
-0.129
(18)
Number
of
flowers
-0.098
(28)
0.512
(4)
0.063
(27)
-0.442
(5)
Length
of
outer
tepal
0.833
(2)
-0.134
(25)
-0.259
(13)
-0.101
(21)
Width
of
outer
tepal
0.688
(5)
-0.369
(8)
-0.300
(11)
-0.166
(14)
Width
streak
on
tepal*
0.472
(17)
0.089
(29)
0.605
(1)
-0.074
(26)
Blotch
length
at
center
0.699
(4)
-0.518
(3)
0.115
(21)
0.099
(24)
Blotch
length
at
margin*
0.560
(14)
-0.535
(2)
-0.077
(26)
-0.001
(30)
Hairs
lower
part
tunic
0.679
(6)
-0.321
(11)
0.363
(5)
0.086
(25)
Hairs
upper
part
tunic
-0.382
(22)
-0.631
(1)
0.330
(8)
-0.074
(26)
Number
of
offset
bulbs
0.353
(23)
0.305
(14)
-0.322
(9)
0.473
(3)
Table
3.
Percentage
of
explained
variation,
and
the
factor
loads
and
the
ranking
in
paren-
theses
of
the
16
main
determining
characters
for
the
first
four
principal
components
in
the
1989
dataset
with
sects.
Eichleres
and
Tulipa.
*
Characters
with
high
factor
loads
exclu-
sively
in
1989
First
PC
18.62%
Second
PC
8.86%
Third
PC
8.60%
Fourth
PC
7.93%
Flowering
date*
0.448
(11)
-0.060
(25)
-0.426
(4)
0.353
(7)
Plant
length*
0.033
(26)
-0.475
(5)
-0.350
(8)
-0.214
(11)
Stem
diameter
-0.837
(1)
-0.239
(11)
-0.096
(21)
-0.089
(23)
Stem
pubescence
-0.507
(9)
0.066
(24)
0.534
(2)
0.133
(19)
Number
of
leaves*
-0.035
(25)
0.203
(14)
0.517
(3)
0.245
(10)
Leaf
undulation
0.314
(17)
-0.129
(20)
-0.338
(9)
0.641
(2)
Leaf
pubescence
-0.631
(4)
-0.029
(27)
0.385
(6)
0.387
(6)
Width
of
second
leaf
-0.602
(5)
-0.623
(2)
0.141
(20)
-0.200
(12)
Number
of
flowers
0.027
(28)
-0.130
(19)
0.633
(1)
-0.005
(28)
Length
of
outer
tepal
-0.815
(2)
-0.266
(9)
-0.059
(25)
-0.092
(22)
Width
of
outer
tepal
-0.747
(3)
-0.398
(7)
-0.086
(23)
0.143
(17)
Tip
outer
tepal
pub.*
-0.557
(7)
0.211
(12)
0.081
(24)
0.489
(3)
Tip
inner
tepal
pub.*
-0.405
(13)
-0.011
(29)
-0.301
(13)
0.664
(1)
Blotch
length
at
center
-0.579
(6)
0.136
(17)
-0.305
(11)
0.063
(24)
Hairs lower
part
tunic
-0.515
(8)
0.466
(6)
-0.392
(5)
-0.147
(16)
Hairs
upper
part
tunic
-0.447
(12)
0.635
(1)
-0.303
(12)
-0.293
(9)
Bulb
tufted
at
top*
-0.317
(16)
0.614
(3)
-0.149
(19)
-0.120
(21)
Number
of
offset
bulbs
0.185
(20)
-0.503
(4)
-0.196
(17)
0.030
(26)
20
L.
W.
D.
VAN
RAAMSDONK
8L
T.
DE
VRIES:
Results
The
statistics
of
the
independent
analyses
of
the
1988
dataset
with
30
characters
and
the
1989
dataset
including
29
characters
are
presented
in
Tables
2
and
3,
respectively.
Every
character
having
a
factor
load
exceeding
0.5
is
included
in
the
tables.
These
characters
are
assumed
to
have
a
major
influence
on
the
direction
of
the
respective
principal
component,
which
is
referred
to
as
a
"high
load".
The
first
and
second
Principal
Component
(PC)
in
the
1988
analysis
showed
a
higher
per-
centage
of
explained
variation
than
the
same
PC's
in
the
1989
analysis.
Up
to
four
PC's
are
needed
in
order
to
take
a
sufficient
part
(approx.
50%)
of
total
variation
into
account.
Of
all
characters
only
two
show
a
high
factor
load
in
more
than
one
PC,
i.e.,
blotch
length
at
center
of
the
tepals
in
the
1988
analysis
(Table
2)
and
width
of
second
leaf
in
the
1989
analysis
(Table
3).
The
first
PC
is
predominantly
determined
by
five
characters,
which
range
among
the
six
highest
factor
loads
in
both
years.
For
the
second
PC
only
the
hairiness
of
the
upper
part
of
the
bulb
tunic
appeared
almost
equally
important
in
both
years.
Several
characters
range
among
those
with
the
highest
factor
loads
for
more
than
one
PC
per
year
indicated
by
the
ordinal
numbers
(Tables
2
and
3).
The
general
distributions
of
the
specimens
in
the
1988
and
1989
analyses
are
shown
in
Figs.
1
and
2,
respectively.
Sect.
Clusianae
takes
a
separated
region
in
the
PCA
diagrams
of
both
years.
The
other
sections
cover
regions
which
overlap
partly.
The
main
part
of
each
section,
however,
possesses
recognizable
parts
of
the
attribute
space.
Some
species
were
found
isolated
from
the
main
part
of
the
respective
sections,
like
T.
praecox
of
sect.
Tulipanum
(Fig.
1)
and
T.
kuschkensis,
T.
lanata,
and
T.
ingens
of
sect.
Eichleres
(Fig.
2).
In
order
to
analyse
the
positions
of
all
other
species
more
in
detail,
quarter
plots
were
made:
Figs.
3,
4,
and
5
show
detailed
parts
of
Fig.
1,
whereas
Figs.
6,
7,
8,
and
9
show
detailed
parts
of
Fig.
2.
For
sect.
Eichleres
a
vertical
border,
indicated
by
a
broken
line,
with
a
value
of
—0.6
at
the
first
PC
is
used
to
distinguish
between
several
species
groups.
For
sect.
Clusianae
two
main
subregions
were
found,
distinguished
by
second
and
third
PC
(Fig.
3):
T.
linifolia,
T.
batalinii,
and
T.
maximowiczii
with
low
val-
ues,
and
T.
aitchisonii
and
T.
clusiana with
high
values.
T.
montana
takes
an
inter-
mediate
position.
Yellow
flowered
specimens,
originally
identified
as
T.
chrysan-
tha,
the
yellow
variety
of
T.
montana
(heavy
cross
signs,
Fig.
3),
appeared
to
be
indistinguishable
from
T.
batalinii.
The
typical
variety
of
T.
clusiana
and
variety
chrysantha
show
a
slight
difference
for
the
third
PC.
The
fourth
PC
appeared
to
be
of
no
importance
for
distinction
between
groups.
All
species
in
sect.
Kolpakowskianae
were
located
in
the
same
part
of
the
vari-
ation
denoted
by
the
first
PC,
i.e.
no
differentiation
along
the
horizontal
axis
is
present
(Fig.
4).
The
two
main
species
of
the
section,
T.
kolpakowskiana
and
T.
ostrowskiana,
cover
separated
regions
when
taking
the
other
PC's
into
account.
T.
tetraphylla
is
indistinguishable
from
T.
ostrowskiana.
T.
iliensis
and
T.
ferganica,
and
T.
behmiana
were
found
in
the
same
regions
as
T.
kolpakowskiana
and
T.
ostrowskiana,
respectively,
except
for
some
differences
at
the
third
PC.
T.
zenai-
dae
and
T
lehmanniana
take
an
intermediate
position.
Several
species
of
sect.
Tulipanum
show
well
separated
regions
in
the
PCA
dia-
grams
(Fig.
5),
especially
T.
praecox.
On
the
other
hand,
T.
systola,
T.
stapfii,
and
T.
ulophylla
were
found
in
the
same
region.
-
2
0
-
subgenus
Tulipa
p.p.
(1988)
2
-
1
-
-1
-
0
0
°3
0
p
0
,
41
ea
pe
01
0
No
00
0
8
00
b
x
.
A
A
0
x
-2
0
A
A
A
14
AA
Tulipa
subg.
Tulipa
21
0
a
cc
N
2
-
LEA
A
X
X•
x
x
x x
X
-1
-
A(x
&
A
A
A
El
AA
9
o
A
A
i
-2
-
0
2
first
pc
Fig.
1.
Principal
component
plot
with
the
first
and
second
and
first
and
third
principal
components
based
on
an
analysis
of
the
1988
dataset
(see
material
and
methods).
Signs:
small
tulip
flowers:
sect.
Clusianae;
cross:
sect.
Kolpakowskianae;
upper
triangles:
sect.
Tulipanum
(black
upper
triangles:
T.
praecox);
grey-shaded
rhombs
and
squares:
sect.
Eichleres
(grey
squares:
T
fosteriana;
grey
rhombs:
T.
vvedenskyi;
lower
triangles:
T.
tschimganica
and
T.
kaufmanniana;
dark
grey
rhombs:
T.
praestans);
wide
tulip
flowers:
sect.
Tulipa.
The
latter
two
sections
were
used
as
reference
to
the
1989
dataset
The
species
of
sect.
Eichleres
with
a
value
of
less
than
—0.6
for
the
first
PC
are
shown
in
Fig.
6.
A
group
consisting
of
T.
tubergeniana,
T
hoogiana,
T.
fosteriana,
T.
eichleri,
and
T.
micheliana
can
be
distinguished.
However,
a
second
group
with
T.
greigii
and
T.
mogoltavica
share
the
same
regions
for
all
PC's.
These
two
groups
can
be
distinguished
by
additional
characters
not
included
in
the
PCA
dataset,
like
00
e
;
ye
°
0
)11
00A
22
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
subgenus
Tulips
p.p.
(1989)
3
2
0
-1
-
00
-2
-
••
••
S[2
7
3
2
-1
-
-2
-
0
0
o
o
0
E
p
O
0
0
00
0
El
0
RoP
1
a
-o
C
0
-
8
V1
0
-3
-1
1
first
pc
Fig.
2.
Principal
component
plot
with
the
first
and
second
and
first
and
third
principal
components
based
on
an
analysis
of
the
1989
dataset
(see
material
and
methods).
Signs:
small
tulip
flowers:
sect.
Clusianae;
cross:
sect.
Kolpakowskianae;
upper
triangles:
sect.
Tulipanum;
rhombs,
squares
and
lower
triangles:
sect.
Eichleres
(the
species
indicated
by
their
first
letter
are:
T.
lanata,
T.
ingens,
T.
kuschkensis,
and
T.
subpraestans;
grey
squares:
T.
eichleres
c.s.
and
T.
greigii
c.s.;
grey
rhombs:
T.
vvedenskyi
c.s.;
lower
triangles:
T.
kaufmanniana
c.s.;
dark
grey
rhombs:
T.
praestans);
wide
tulip
flowers:
sect.
Tulipa.
The
first
three
sections
were
used
as
reference
to
the
1988
dataset
Tulipa
subg.
Tulipa
23
anthocyanidin
stripes
on
the
upper
surface
of
the
leaves
and
shape
and
colour
intensity
of
the
blotch.
Several
groups
of
species
of
sect.
Eichleres
showed
values
higher
than
—0.5
at
the
first
PC.
The
species
under
consideration
excluding
those
of
sect.
Spiranthera
(Fig.
7)
possess
different
areas,
although
overlap
exists
with
respect
to
the
second
and
fourth
PC.
These
areas
are
referred
to
as
species
regions
with
intermediate
overlapping
zones.
These
regions
allow
comparison
with
the
species
of
sect.
Spi-
ranthera,
located
in
the
same
part
of
the
total
PC
diagram.
T.
praestans
and
T.
sos-
novskyi
are
clearly
separated.
T.
vvedenskyi
shares
the
same
region
with
T.
alber-
tii
and
T.
butkovii.
The
species
originally
considered
to
belong
to
sect.
Spiranthe-
ra
are
shown
in
Fig.
8,
with
an
indication
of
the
species
regions
as
indicated
in
Fig.
7.
These
species
predominantly
share
region
2B
with
T.
vvedenskyi,
T.
alber-
tii,
T.
butkovii,
and
T.
praestans,
region
3B
with
T
vvedenskyi,
T.
albertii,
and
T.
butkovii,
and
region
4A
with
T.
sosnovskyi.
T.
kaufmanniana
was
hardly
found
in
the
overlapping
zones
between
regions,
whereas
T.
tschimganica,
T.
anadroma,
and
T.
dubia
show
similarities
to
specimens
in
adjacent
regions.
In
sect.
Tulipa,
T.
rhodopea,
and
T.
hungarica
are
located
at
the
same
regions
in
all
diagrams
(Fig.
9).
Two
specimens
of
T.
rhodopea
show
a
deviating
habitus
with
respect
to
the
characters
as
loaded
on
the
first
PC.
T.
suaveolens
is
located
outside
the
region
in
the
diagram
covered
by
T.
rhodopea
in
all
but
one
cases.
T.
armena
and
the
so-called
Neotulipae
T
didieri,
T.
planifolia,
T.
marjolettii,
and
T.
grengiolensis,
occupy
two
different
regions
with
hardly
any
overlap
in
the
PCA
diagrams.
The
forms
of
T.
gesneriana
show
considerable
variation,
greatly
outside
the
regions
of
the
species
discussed.
In
general
the
characters
show
high
correlation
coefficients
(R
>
0.85)
between
the
years,
which
indicates
high
genetic
control
and
reliability.
Exceptions
are
plant
length
(R
=
0.705),
leaf
undulation
(R
=
0.786),
number
of
flowers
(R
=
0.305)
and
bulb
tunic
colour
(R
=
0.525).
All
values
in
the
1988
dataset
of
the
characters
meas-
ured
on
a
continuous
scale
were
transformed
to
the
1989
data
ranges
based
on
lin-
ear
regression
of
the
15
reference
populations
between
the
years.
Of
these
charac-
ters,
only
number
of
flowers
was
not
excluded
from
the
combined
dataset,
since
the
low
correlation
coefficient
was
exclusively
due
to
the
variation
in
T
praestans.
Table
4.
Percentage
of
explained
variation,
and
the
factor
loads
and
the
relative
ranking
in
parentheses
of
the
eight
main
determining
characters
for
the
first
two
canonical
variates
in
the
total
dataset
of
subg.
Tulipa
First
CV
40.83%
Second
CV
16.38%
Stem
pubescence
1.858
(2)
—0.133
(14)
Leaf
pubescence
1.245
(4)
1.766
(1)
Teal
outside
pubescent
0.837
(7)
1.364
(2)
Width
streak
blotch-tip
0.007
(21)
0.525
(5)
Hairy
coronet
at
base
2.110
(1)
—0.448
(6)
Hairs
lower
part
tunic
1.455
(3)
0.081
(15)
Hairs
upper
part
tunic
—1.230
(5)
0.646
(3)
Bulb
tufted
at
top
—1.162
(6)
0.637
(4)
24
L.
W.
D.
VAN
RAAMSDONK
8L
T.
DE
VRIES:
Table
5.
Global
amount
of
heterochromatin
after
C-banding
of
Tulipa
species.
The
numbers
refer
to
the
accession
Section
Heterochromatin
none
or
low
moderate
high
Clusianae
T
batalinii:
64181
70597
T.
limfolia:
67261
70596
Kolpakowskianae
T.
kolpakowskiana:
68596
74228
T.
ostrowskiana:
68055
70600
78120
T.
tetraphylla:
65302
67368
Tulipanum
Tulipanum
T.
agenensis:
77310
T.
agenensis:
84259
T.
Julia:
72119
84260
T.
stapfii:
71341
T.
systola:
79154
Aureo-fasciatae
T.
praecox:
67741
77101
Eichleres
Lana
tae
T.
ingens:
74429
Eichleres
T.
eichleri:
67234
T.
hoogiana:
82116
T.
fosteriana:
71324A
Vinistriatae
T.
greigii:
78122
Undulatae
T.
vvedenskyi:
81120
T.
vvedenskyi:
68087
Multiflorae
T.
praestans:
70626
77266
Spiranthera
T.
kaufmanniana:
65301
T.
kaufmanniana:
65252
T.
tschimganica:
76123
68014
T.
amblyophylla:
74111
Tulipa
T.
armena:
74223
T.
hungarica:
66066
T.
galatica:
84266
T.
planifolia:
67737
76181
T.
rhodopea:
75108
T.
suaveolens:
65038
T.
marjolettii:
66040
73120
T.
grengiolensis:
67738
The
combined
dataset
with
measurements
of
1989
and
those
of
1988
recalcu-
lated
to
the
1989
data
ranges
and
containing
24
characters,
was
subjected
to
Canonical
variate
analysis.
The
percentage
of
explained
variation
of
the
first
and
second
canonical
variate
(CV)
and
the
8
characters
with
the
highest
factor
loads
are
presented
in
Table
4.
All
most
important
characters
showed
aspects
of
the
Tulipa
subg.
Tulipa
25
degree
of
hairiness.
The
characters
with
the
highest
loads
on
the
second
CV
were
also
included
in
the
seven
most
important
characters
of
the
first
CV.
A
CVA
diagram
with
species
groups
plotted
against
the
first
and
second
CV
is
shown
in
Fig.
10.
Sect.
Clusianae
is
separated
from
the
other
sections
as
after
PCA.
T.
kuschkensis
(group
2)
is
located
among
the
species
of
sect.
Tulipanum.
The
species
groups
with
T.
lanata
c.s.,
T.
eichleri
c.s.,
and
T.
greigii
c.s.,
groups
1,
3,
and
4,
respectively,
with
values
for
the
first
PC
lower
than
—0.5
are
also
separ-
ated
from
the
rest
of
sect.
Eichleres
in
Fig.
10.
The pairs
T.
eichleri
c.s.
and
T.
grei-
gii
c.s.
(groups
3
and
4),
T.
armena
c.s.
and
the
Neotulipae
species
(groups
A
and
C),
T.
clusiana
and
T.
aitchisonii
(groups
D
and
E),
and
T.
julia
and
T.
aleppensis
(groups
J
and
K)
are
hardly
distinguishable
in
the
CVA
plot.
T.
praestans
shows
a
tendency
towards
sect.
Tulipanum.
A
C-banding
pattern
with
a
high
number
of
bands
is
shown
in
Fig.
11.
The
pat-
terns
appeared
to
be
unique
for
each
species
and
a
relation
between
band
positions
in
different
species
could
not
be
established.
The
global
amount
of
heterochroma-
tin
is
presented
in
Table
5.
Hardly
any
heterochromatin
(less
than
2%)
was
found
in
sects.
Kolpakowskianae
and
Eichleres,
while
high
levels
(more
than
20%)
were
present
in
some
species
of
sect.
Tulipanum
and
Tulipa.
Accessions
showing
no
heterochromatin
were
tested
for
several
subsequent
years.
Some
accessions
additional
to
those
used
for
multivariate
analysis
have
been
identified
as
indicated
in
Table
1
and
were
used
to
complete
the
view
on
and
the
documentation
of
the
species
with
respect
to
morphology,
chromosome
number
and
distribution.
Discussion
Genus
typification.
LINNAEUS
(1753)
described
two
species:
T.
gesneriana
(subg.
Leiostemones)
and
T
sylvestris
(subg.
Eriostemones).
A
third
one
from
Ethiopia,
T.
breyniana,
is
now
placed
in
the
genus
Baeometra.
Presumably
T.
gesneriana
fits
best
LINNAEUS'
concept
of
the
genus,
since
specimens
of
T.
gesneriana
were
avail-
able
in
the
CLIFFORD
Herbarium,
where
T.
sylvestris
is
not
represented
(UNNAEus,
1737
a).
So,
T.
gesneriana
was
the
first
species
known
to
LINNAEUS
and
probably
the
only
one
at
the
time
he
wrote
the
first
edition
of
Genera
Plantarum
(1737
b).
Moreover,
T
gesneriana
is
suited
better
to
the
general
idea
of
the
typical
pheno-
type
of
a
tulip.
T.
gesneriana
is
designated
the
type
of
the
genus,
and
subg.
Leiostemones
is
therefore
formally
named
subg.
Tulipa.
This
choice
has
been
made
by
DASGUPTA
&
DEB
(1985)
and
was
followed
by
JARVIS
&
al.
(1992,
1993).
Subgeneric
division.
Neither
BOISSIER
(1882)
nor
HALL
(1940)
gave
an
indica-
tion
of
the
level
of
the
two
main
groups
in
the
genus,
although
HALL
(1940)
subdi-
vided
the
Leiostemones
into
five
subsections
and
the
Eriostemones
into
three
sec-
tions.
Both
main
groups
were
considered
sections
by
STORK
(1984)
and
MARAIS
(1984).
The
current
subdivision
in
sects.
Eriostemones
and
Leiostemones
is
now
well
documented
by
differences
in
morphology
(HALL
1940,
STORK
1984,
VAN
RAAMSDONK
&
DE
VRIES
1992),
flower
pigment
composition
(NIEUWHOF
&
al.
1990)
and
absence
of
crossability
relationships
(VAN
EIJK
&
al.
1991).
On
account
of
these
differences
the
sections
are
raised
to
the
level
of
subgenera.
In
addition,
the
group-
ings
described
as
subsections
(HALL
1940,
STORK
1984)
are
raised
to
the
level
of
sections.
Formal
descriptions
are
given
in
the
chapter
taxonomic
enumeration.
26
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
The
five
sections
of
subg.
Tulipa
show
overlapping
regions
in
at
least
two
of
the
four
PC's
used
in
this
study
(Figs.
1
and
2),
except
for
sect.
Clusianae
which
is
clearly
separated
morphologically.
The
existing
division
into
sections
(HALL
1940:
subsections)
is
maintained
because
of
the
general
differences
in
morphology
between
the
sections
(Figs.
1,
2,
and
10)
and
of
the
different
geographical
distri-
butions
(Hoot
1973),
although
some
species
are
replaced
from
one
section
to
another.
For
priority
reasons,
the
name
Scabriscapae
(BAKER
1874)
should
be
used
instead
of
one
of
the
names
used
by
HALL
(1940).
BAKER
(l.c.)
included
species
now
classified
in
sects.
Kolpakowskianae
,
Eichleres,
and
Tulipa,
and
the
description
of
Scabriscapae
covers
(part
of)
these
sections
as
well.
BAKER
(1883)
did
not
main-
tain
a
section
or
subsection
under
the
name
Scabriscapae
and
he
classified
the
species
included
in
Scabriscapae
(BAKER
1874)
in
the
large
sect.
Gesnerianae
(BAKER
1883).
So,
the
meaning
of
this
name
is
unclear
and
it
must
be
stated
a
nomen
nudum.
Character
distribution
and
species
delimitation.
The
high
correlation
coef-
ficients
of
most
characters
between
years
point
to
a
high
repeatability.
The
genetic
component
appeared
to
be
dominant
in
the
phenotype.
Nevertheless,
the
geno-
type-environment
interaction
is
unknown
and
a
straightforward
conclusion
of
high
heritability
figures
cannot
be
drawn.
The
absolute
date
of
flowering
may
vary
between
years
depending
on
temperature
and
rainfall
but
the
species
appeared
to
flower
in
almost
identical
order
in
the
years
investigated,
which
caused
the
high
correlation coefficient.
Among
the
characters
used
for
the
several
PC
and
CV
analyses
the
characters
concerning
hairiness
or
pubescence,
especially
of
the
bulb
tunic
range
as
most
important
(Tables
2,
3,
and
4).
This
importance
supports
the
recognition
of
sects.
Clusianae
and
Tulipanum
of
which
the
species
show
the
characters
bulbs
tufted
with
hairs
and
upper
parts
of
the
bulb
tunics
hairy,
and
bulb
tunics
entirely
wool-
ly,
respectively.
A
correlation
with
geographic
distribution
may
occur
since
the
species
of
these
sections
predominantly
occur
south
of
major
mountain
ranges,
i.e.,
sect.
Clusianae
south
of
the
Pamir
Alai
and
Himalaya
and
sect.
Tulipanum
south
of
the
Swiss,
Austrian
and
Julian
Alps,
Caucasus
and
Kopeth
Dagh
(Hoot
1973).
Any
ecological
relevance
or
selective
advantage
cannot
be
indicated.
Woolly
bulb
tunics
could
be
considered
to
be
advantageous
in
hardish
climates,
but
with-
in
sect.
Biflores
(subg.
Eriostemones)
species
with
either
woolly
or
glabrous
bulb
tunics
have
been
found
in
Central
Asian
mountain
regions
as
well
as
in
South
Rus-
sia
and
the
Balkan
peninsula
(VAN
RAAMSDONK
&
DE
VRIES
1992).
Sect.
Tulipanum
shares
its
geographic
distribution
area
with
species
of
sect.
Tulipa
possessing
almost
glabrous
bulb
tunics.
Flower
colour
within
species
may
differ
in
two
respects.
At
first,
blotch
(the
basal
parts
of
the
tepals,
usually
darker
than
main
flower
colour)
and
blotch
mar-
gin
(usually
yellow)
may
show
difference
in
size
as
well
as
in
intensity.
Secondly,
within
some
species
anthocyanidins
are
lacking
in
certain
accessions
resulting
in
yellow
or
very
light
colours
(NIEuwHoF
&
al.
1990).
After
selection
experiments
in
accessions
obtained
from
natural
provenances
(EIKELBoom,
CPRO-DLO,
unpubl.)
as
well
as
mutation
experiments
with
radiation
(BROERTJES
&
VAN
HARTEN
1988)
it
was
shown
that
blotch
margin
and
flower
colour
can
easily
be
influenced.
In
sev-
a
_c
t
a
0
-
o
0
Tulipa
subg.
Tulipa
27
section
Clusianae
2
a
1
-
0
-
section
Kolpakowskianae
1
-
0
-1
-
+
+
+
o
)
,!
051
+
+
p
p
A
A
-1
-
* t
40
A
-2
2
2
2
1
++,v
Vo'l
5 t
4+
+
V
EV
+
.
1
0
-
B
X
C
X
a
-
2
0
-1
-
:E
o
q
-1
-
-2
1
-2
+
ig
0
VCJ",'•
p-PP
,
p+
6
X
+
0
+
-1
-
Sc
0
El
\x
i
,
e
>14
-2
-2
0
0
first
pc
first
pc
Fig.3
Fig.
4
Fig.
3.
Part
of
a
principal
component
plot
with
the
first
and
second,
first
and
third,
and
first
and
fourth
principal
components
based
on
an
analysis
of
the
1988
dataset
(see
material
and
methods).
Signs:
plus:
T.
aitchisonii;
lower
triangle:
T.
clusiana
var.
chrysantha;
upper
triangle:
T.
clusiana
var.
clusiana;
grey
squares:
T.
montana;
cross:
T.
linifolia
(incl.
T.
batalinii
and
T
maximowiczii);
heavy
cross:
T
montana
yellow
variety
Fig.
4.
Part
of
a
principal
component
plot
with
the
first
and
second,
first
and
third,
and
first
and
fourth
principal
components
based
on
an
analysis
of
the
1988
dataset
(see
material
and
methods). Signs:
plus:
T
kolpakowskiana;
grey
rhombs:
T.
iliensis;
upper
triangles:
T.
ferganica;
heavy
cross:
T.
zenaidae;
heavy
plus:
T.
lehmanniana;
open
squares:
T
behmi-
ana;
grey
squares:
T.
tetraphylla;
black
squares:
T.
ostrowskiana
28
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
eral
species
a
yellow
blotch
margin
is
either
present
or
absent,
i.e.,
in
T
stapfii,
T.
lanata,
T.
ingens,
T.
hoogiana,
and
T.
fosteriana.
Species
described
as
closely
related
(HALL
1940,
BOCHANTZEVA
1982,
STORK
1984)
may
differ
exclusively
in
flower
colour-like
T.
clusiana
and
T.
stellata,
T.
linifolia
and
T.
batalinii,
and
T.
armena
and
T.
galatica.
Species
are
described
as
formas
when
the
flower
colour
is
the
only
distinguishing
feature.
In
several
cases
species
show
slight
morphological
differences
but
differ
greatly
in
their
geographic
distribution.
In
the
present
study
such
species
pairs
are
treated
on
the
level
of
subspecies
when
allopatric
or
as
varieties
when
partly
sympatric.
New
combinations
are
described
in
the
chapter
taxonomic
enumeration.
C-banding.
Data
on
chromosome
C-banding
were
reported
by
BLAKEY
&
VOSA
(1982),
predominantly
using
material
from
the
CPRO-DLO
collection
(formerly
IVT).
Hardly
any
heterochromatin
was
found
in
sect.
Kolpakowskianae
and
most
species
of
sect.
Eichleres,
although
T.
fosteriana
and
T.
greigii
showed
moderate
amounts
(BLAKEY
&
VOSA
1982),
which
is
contrary
to
our
present
results
(Table
5).
In
sect.
Tulipa
a
moderate
number
of
C-bands
were
organized
around
the
centro-
mere
(BLAKEY
&
VOSA
1982).
This
phenomenon
was
only
partly
found
in
T.
galati-
ca,
while
the
minor
chromosomes
were
significantly
shorter
(Fig.
11)
than
reported
by
BLAKEY
&
VOSA
(1982).
Moderate
banding
in
sect.
Clusianae
and
moderate
to
strong
banding
in
sect.
Tulipanum
(BLAKEY
&
VOSA
1982)
is
in
concordance
with
the
present
results
(Table
5).
The
statement
of
LoIDL
(1983),
that
a
high
amount
of
heterochromatin
is
an
advanced
characteristic,
should
imply
that
sect.
Kolpakows-
kianae
is
the
most
primitive
and
that
the
most
advanced
species
were
found
in
sects.
Tulipanum
and
Tulipa.
Sect.
Clusianae.
Species
of
sect.
Clusianae
are
found
south
of
the
mountain
ranges
Kopeth
Dagh,
Pamir
Alai
and
western
Himalayas,
the
T.
linifolia
complex
originally
assigned
to
be
endemic
in
the
Pamir
Alai
(Hoot
1973).
T.
clusiana
f.
clusiana
shows
white
or
cream
flowers
of
which
the
outer
tepals
have
a
broad
red
or
crimson
band
at
the
dorsal
side.
The
blotch
is
purple
(HALL
1940).
T.
stellata
has
a
yellow
blotch,
while
its
form
chrysantha
is
completely
yellow.
One
collected
sample
(GWH
638)
was
provisionally
identified
as
T.
clusiana
yellow
variety
(HEWER
1974)
but
it
appeared
to
consist
of
individuals
belonging
to
T.
batalinii
as
well.
T.
clusiana,
T.
stellata,
and
"T.
stellata
chrysantha"
were
considered
to
be
tetraploid
or
pentaploid
(HALL
1938,
1940).
Plants
collected
at
the
end
of
the
19
th
century
with
flowers
like
the
typical
T.
clusiana
proved
to
be
diploids
and
were
assigned
to
T.
aitchisonii
(HALL
1938).
Later
also
completely
yellow
forms
on
the
diploid
level
were
collected
and
described
as
T.
aitchisonii
subsp.
cashmeriana
(HALL
1938,
1940).
T.
aitchisonii
is
considered
synonymous
to
T.
clusiana
(SEALY
1948),
since the
variation
in
morphology
(Fig.
3)
and
flower colour
appeared
to
be
identical.
The
four
formas
as
described
in
taxonomic
enumeration
therefore
con-
sist
of
diploids
as
well
as
polyploids.
The
species
T.
fernandezii
and
T.
porphyreo-
chrysantha
of
the
T.
clusiana
aggregate
(BLATTER
&
FERNANDEZ
1934)
both
possess
yellow
flowers
without
a
blotch
and
with
a
crimson
dorsal
band
on
the
tepals.
The
type
of
T.
porphyreo-chrysantha
is
indicated
by
BLATTER
&
FERNANDEZ
(1934)
as
syntype
of
T.
fernandezii.
The
cited
herbarium
sheets
were
not
found;
they
should
be
present
in
the
Bombay
BLATTER
herbarium
(DASGUPTA
&
DEB
1985),
but
we
were
not
able
to
locate
them.
Both
species
have
been
treated
as
nomina
dubia
within
T.
Tulipa
subg.
Tulipa
29
clusiana
(DASGUPTA
&
DEB
1985).
The
name
T.
chrysantha
Boiss.
was
originally
published
to
describe
plants
which
are
currently
regarded
as
yellow
variants
of
T.
montana
(q.v.).
However,
BAKER
(1874)
included
apparently
plants
belonging
to
the
T.
clusiana
aggregate.
So,
several
combinations
with
the
epitheton
"chrysan-
tha"
have
been
used
incorrectly,
T.
clusiana
var.
chrysantha
(HALL)
SEALY
amongst
them
(HALL
1940,
SEALY
1948).
We
now
conclude
from
the
discussed
nomenclatural
confusions
that
the
names
published
by
BLATTER
and
the
combination
of
SEALY
are
invalid.
T.
batalinii
and
T.
maximowiczii
were
described
as
being
very
closely
related
to
T.
linifolia,
the
latter
with
prostrate
leaves
and
red
flowers
(HALL
1940).
They
are
different
from
T
linifolia
by
yellow
flowers
and
erect
leaves,
respectively.
T
linifolia,
T.
maximowiczii,
and
T.
batalinii
are
treated
as
synonymous
(DYKES
1925);
the
name
T
linifolia
has
priority.
The
name
T.
montana
has
been
applied
to
two
different
taxa;
REGEL
(1873)
used
the
name
twice
in
two
different
species
groups.
LINDLEY
(1827)
assigned
this
name
to
a
species
which
should
be
related
to
T.
linifolia
according
to
DYKES
(1925),
HALL
(1940)
and
SEALY
(1963).
This
species
was
later
described
as
T.
wilsoniana
(HooG
1902).
Later
the
name
T.
montana
was
commonly
used
for
a
species
of
sect.
Tulipanum
with
T.
systola
as
correct
name
(BAKER
1883,
HALL
1940,
SEALY
1963,
MOUTERDE
1966).
A
yellow
variant
of
T.
montana
was
described
as
T
chrysantha
(BoissIER
1882,
SEALY
1948).
All
acces-
sions
used
in
the
present
study
identified
as
T.
montana
appeared
to
be
part
of
the
variation
of
sect.
Clusianae
(Fig.
3).
The
red
flowered
variants
are
intermediate
between
T.
linifolia
and
T.
clusiana
and
designated
as
species.
The
yellow
vari-
ants,
however,
were
indistinguishable
from
accessions
originally
identified
as
T.
batalinii.
As
a
result
the
yellow
flowering
variety
of
T.
linifolia
has
been
given
the
name
var.
chrysantha.
Some
accessions
of
T.
clusiana
and
T.
linifolia
var.
chrysan-
tha
were
found
west
of
the
known
distribution
area
(Table
1;
HALL
1940,
HooG
1973).
These
two
taxa
from
Kashmir
and
the
Pamir
Alai
mountains,
respectively,
apparently
extend
their
distribution
to
Iran.
Sect.
Kolpakowskianae.
The
distribution
of
the
species
of
sect.
Kolpakowskia-
nae
is
predominantly
in
the
Tien
Shan
mountain
range;
some
of
these
species
extend
into
western
China.
T.
ferganica
is
located
in
the
mountains
adjacent
to
the
Fergana
Basin
and
the
distribution
of
T.
lehmanniana
is
extended
to
the
Kyzyl
Kum
area,
Hindu
Kush
and
Kopeth
Dagh
(HooG
1973).
T.
iliensis,
T.
kolpakows-
kiana,
and
T.
ferganica
share
almost
the same
region
in
the
principal
component
plots
(Fig.
4).
Only
on
the
third
PC
some
differences
were
found
in
this
group
(Fig.
4
b).
The
same
applies
to
the
group
with
the
species
T.
tetraphylla,
T.
behmi-
ana,
and
T.
ostrowskiana.
Both
groups
are
considered
to
be
monospecific.
Accord-
ing
to
priority
rules
these
species
are
designated
as
T.
iliensis,
primarily
consisting
of
diploids,
and
T.
tetraphylla,
exclusively
consisting
of
tetraploids
(HALL,
1940,
BOCHANTZEVA
1982,
KROON
&
JONGERIUS
1986).
T.
ferganica
and
T.
ostrowskiana,
respectively,
are
treated
as
subspecies
because
of
their
different
geographic
distri-
bution
(HooG
1973).
Both
species
have
yellow
flowers
tinged
with
red
or
lilac
as
found
in
sect.
Clusianae.
In
T.
ostrowskiana
accessions
with
completely
red
flow-
ers
were
found.
T.
lehmanniana
and
T.
zenaidae
from
the
lower
slopes
of
the
Ala-
tau
valley
(BOCHANTZEVA
1982)
remain
as
separate
species,
having
a
distinctive
brown
blotch.
Sect.
Tulipanum
(Oculus-solis
sensu
HALL).
The
name
Tulipanum
was
origi-
2
-
-2
-
V
V
7
.
7
1>
to
X
0
-
0
1
1
X0(
A
X
V
A
F
V
V
0
-1
-
0
V
V
V
0
-H-
++
A
-1
-
+
>
4
NO
X
0
30
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
section
Tulipanum
0
4
first
pc
Fig.
5.
Part
of
a
principal
component
plot
with
the
first
and
second,
first
and
third,
and
first
and
fourth
principal
components
based
on
an
analysis
of
the
1988
dataset
(see
material
and
methods).
Signs:
rhombs:
T.
systola;
plus:
T
stapfii;
heavy
plus:
T.
ulophylla;
lower
triangles:
T.
oculus-solis;
cross:
T.
julia;
black
upper
triangles:
T.
aleppensis;
black
squares:
T.
praecox;
heavy
cross:
hybrid
between
T.
stapfii
and
T
praecox
nally
used
by
DE
REBOUL
(1847)
to
indicate
a
group
of
species
with
bulb
tunics
woolly
at
the
inside,
containing
T.
praecox,
T.
oculus-solis,
T.
clusiana,
T.
maleo-
lens
and
an
undocumented
species
name.
Later
sect.
Tulipanum
was
considered
to
consist
of
T
tubergeniana,
T.
kuschkensis,
T.
hoogiana,
and
T.
julia
(VVEDENSKY
1935);
all
these
species
appeared
to
be
described
later
than
1847.
The
absence
of
the
species
named
by
DE
REBOUL
(1847)
is
due
to
the
fact
that
they
occur
outside,
1
0
0
'7
0
A
.•
_1
-
a)
CD
x
Tulipa
subg.
Tulipa
section
Eichleres
p.p.
1
31
x x
x
X
X
X›
)
,
x
-2
2
1
0 0
x x
mix
12
o
v
+
vc,
-
1
N S
V'
'7/
-2
-3
first
pc
Fig.
6.
Part
of
a
principal
compo-
nent
plot
with
the
first
and
second,
first
and
third,
and
first
and
fourth
principal
components
based
on
an
analysis
of
the
1989
dataset
(see
material
and
methods).
Signs:
upper
triangles:
T.
hoogiana;
rhombs:
T.
tubergeniana;
lower
triangles:
T.
fosteriana;
cross:
T.
eichleri;
heavy
cross:
T.
micheliana;
black
squares:
T.
greigii;
plus:
T.
mogoltavica
i.e.,
south
of
the
region
covered
by
Flora
of
the
USSR.
HALL
(1940)
treated
the
species
considered
to
belong
to
sect.
Tulipanum
as
a
special
group
within
subsect.
Oculus-solis.
BOCHANTZEVA
(1982)
added
a
fifth
species
to
VVEDENSKY'S
(1935)
enumeration:
T
subquinquefolia
VVED.
The
lack
of
relationship
between
T.
clusiana
at
one
hand,
and
T.
praecox
and
T
oculus-solis
at
the
other
is
obvious.
Therefore,
the
sect.
usually
indicated
as
Oculus-solis
(Eriobulbi/BAKER)
deserves
the
name
Tulipanum
for
priority
reasons.
The
representatives
of
this
section
occur
in
central
Europe,
Asia
Minor,
Lebanon,
Syria,
Caucasus,
Iran,
Iraq,
and
Afghanistan
(Hoot
1973).
The
species
of
this
group
resemble
species
of
sect.
Tulipa
(HALL
1940)
and
2
X
x
X
00
°
O
O
-1
-
-2
-
v
X
X
x
X
X
X
G
0
*
t
*
♦0
*
*
e
. *
4
.
O
a_
region
4A
region
4B
region
3A
region
3B
region
3C
x
>0(
.
°
X
X
X
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
32
section
Eichleres
p.p.
first
pc
region
2A
region
2B
Fig.
7.
Part
of
a
principal
component
plot
with
the
first
and
second,
first
and
third,
and
first
and
fourth
principal
components
based
on
an
analysis
of
the
1989
dataset
(see
material
and
methods).
Signs:
cross:
T.
vvedenskyi;
lower
triangles:
T.
albertii;
upper
triangles:
T.
butkovii;
grey
rhombs:
T.
sosnovskyi;
plus:
T.
praestans
Tulipa
subg.
Tulipa
33
some
species
of
sect.
Eichleres
(Fig.
2)
with
respect
to
foliage
and
flower
charac-
teristics,
but
differ
because
of
the
long,
interwoven
hairs
at
the
inside
of
the
bulb
tunics
(HALL
1940).
The
name
T.
agenensis
was
published
in
February
1804
by
DE
CANDOLLE
in
REDourt
(1804).
Shortly afterwards
SAINT
AMANS
(1804)
discussed
the
name
T.
oculus-solis
for
a
species
which
appeared
to
be
identical
to
that
of
DE
CAN-
DOLLE,
but
SAINT
AMANS
(1804)
did
not
add
a
description.
Therefore,
the
correct
name
of
this
species
is
T.
agenensis
(MARAis
1980).
T
systola
and
T.
stapfii
resem-
ble
each
other
very
closely
and
are
treated
as
conspecific.
The
specific
status
of
T.
aleppensis
is
doubtful
(Figs.
5
and
10).
It
may
well
be
a
triploid
derivative
of
T.
julia,
although
diploid
forms
were
reported
(KRooN
&
JONGERIUS
1986).
T.
stapfii
is
treated
including
T.
ulophylla.
Sect.
Eichleres.
In
sect.
Eichleres,
the
largest
of
the
subg.
Tulipa,
large
differ-
ences
with
respect
to
geographic
distribution
(HooG
1973)
and
morphology
(Figs.
2,
6-8)
were
found.
T.
kuschkensis
with
woolly
bulb
tunics
is
placed
in
sect.
Tulipanum
by
HALL
(1940)
and
HEWER
(1974),
which
is
confirmed
by
the
present
results
(Fig.
10).
T.
ingens
and
T.
lanata
(Pamir
Alay
and
Eastern
Kopeth
Dagh;
HooG
1973),
and
T.
subpraestans
(Pamir
Alay;
HooG
1973)
are
separated
from
the
main
part
of
the
section
(Fig.
2),
and
placed
in
two
series.
The
main
part
of
species
of
sect.
Eichleres
consists
of
several
groups
of
closely
related
species.
Two
groups
with
values
lower
than
—0.6
for
the
first
PC
sharing
the
same
region
for
all
four
PC's
(Fig.
6),
are
separated
geographically:
T.
fosteri-
ana,
T.
hoogiana,
T
tubergeniana,
and
T
eichleri
were
found
in
the
Pamir
Alay
and
the
mountain
ranges
west
of
it,
whereas
T.
greigii
and
T.
mogoltavica
occur
in
montane
Kyzyl-Kum
and
Tien
Shan,
north
of
the
geographic
area
of
the
first
group.
Both
groups
are
described
as
series.
T.
tubergeniana
and
T.
hoogiana
differ
in
area
of
distribution:
T.
tubergeniana
is
endemic
in
the
western
Pamir-Alai
mountain
range
(Bukhara
and
environs),
while
T.
hoogiana
occurs
in
montane
Turkmenistan
extending
to
NE
Iran.
Both
distribution
ranges
are
divided
by
the
river
Amu
Darya
(HooG
1973).
T
hoogiana
is
treated
as
synonymous
to
T.
tuber-
geniana,
as
indicated
by
HALL
(1940)
and
BOCHANTZEVA
(1982),
for
the
absence
of
clear
morphological
differences
(Fig.
6).
The
close
relationship
of
T.
micheliana
from
the
Pamir
Alai
(HooG
1973)
with
T.
eichleri
(HALL
1940)
is
confirmed
and
they
are
treated
as
conspecific.
Similarly,
in
the
region
with
scores
for
the
first
PC
higher
than
—0.6
several
groups
of
closely
related
species
were
found
(Figs.
7
and
8).
T.
sosnovskyi
(Cau-
casus),
T.
albertii,
T.
vvedenskyi
and
T.
butkovii
both
treated
as
conspecific
(Tien
Shan),
and
T.
praestans
(Pamir
Alai)
are
separated
by
the
third
PC
(Fig.
7
b).
Because
of
morphological
(HALL
1940
and
present
study)
and
geographical
differ-
ences
(HooG
1973)
these
groups
are
treated
as
series.
The
species
T.
kaufmanniana,
T.
dubia,
and
T.
tschimganica,
endemic
in
west-
ern
Tien
Shan
(HooG
1973)
share
the
geographic
area
with
T.
albertii.
T.
anadro-
ma
is
treated
as
conspecific
with
T.
tschimganica,
and
T.
amblyophylla
with
T.
kaufmanniana.
Red
flowering
accessions
have
been
assigned
to
T.
dubia.
The
group
of
species
related
to
T.
kaufmanniana
has
been
described
as
sect.
Spiranthe-
ra
(VVEDENSKYI
1935,
BOCHANTZEVA
1982).
In
each
principal
component
plot
of
the
1989
analysis
different
species
combine
in
the
same
region
with
the
species
of
sect.
Spiranthera
(Figs.
7
and
8),
but
these
species
are
separated
after
the
1988
0
-
o
S
o
o
~
-2
-
°
34
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
section
Eichleres
p.p.
(Spiranthera)
region
4A
region
4B
1
0
-1
o
I:12
0
0
l
a
0
EL-i0
.•
region
3B
region
3C
1
region
2A
o
o
A
-1
-
op
00
0
region
2B
-1
1
first
pc
Fig.
8.
Part
of
a
principal
component
plot
with
the
first
and
second,
first
and
third,
and
first
and
fourth
principal
components
based
on
an
analysis
of
the
1989
dataset
(see
material
and
methods).
Signs:
squares:
T.
kaufmanniana;
grey
rhombs:
T.
dubia;
black
upper
trian-
gles:
T.
tschimganica;
grey
upper
triangles:
T.
anadroma
analysis
(Fig.
1).
This
section
can
easily
be
recognized
by
means
of
some
charac-
teristics,
which
were
not
included
in
the
presently
used
set
of
characters
for
mul-
tivariate
analyses,
like
leaf
colour,
flower
shape,
and
a
red
spot
at
the
inner
side
of
the
tepals
at
the
place
where
the
margin
of
the
blotch
should
be
located
(HALL
1940).
Besides
totally
red
flowers
in
T.
dubia
as
found
in
all
other
species
of
sect.
Eichleres,
cream
or
yellow
flowers
tinged
with
red
on
the
dorsal
side
occur,
com-
parable
to
the
flower colour
in
sect.
Clusianae
and
Kolpakowskianae
(BOCHANTZE-
VA
1982).
The
accessions
of
T.
kaufmanniana
and
T.
amblyophylla
show
moderate
+
0+
e
v3
v
,
A-
6,
v
v
AA
g
6
0
E3
4AT
-
0--d]
V
t.
31(
0
*
6
-
31E
-e
1
0
U
0.
-1
-
6
-2
-
1
o
0
0
C
-1
-
-2
-
1
0
0
0
U
-1
-2
-3
1
first
pc
Tulipa
subg.
Tulipa
35
section
Tulipa
Fig.
9.
Part
of
a
principal
component
plot
with
the
first
and
second,
first
and
third,
and
first
and
fourth
principal
components
based
on
an
analysis
of
the
1989
dataset
(see
material
and
meth-
ods).
Signs:
dotted
squares:
T.
armena;
grey
rhombs:
T.
hungarica;
plus:
T.
rhodopea;
black
squares:
T.
suaveo-
lens;
upper
triangles:
T.
marjolettii
and
T.
grengiolensis;
lower
triangles:
T.
didieri
and
T.
planifolia;
stars:
T.
ges-
neriana
levels
of
heterochromatin
in
contrast
to
most
other
species
in
sect.
Eichleres
(Table
5).
However,
a
separate
section
can
not
be
maintained
and
the
section
is
therefore
treated
as
series.
Sect.
Tulipa
(Gesnerianae
sensu
HALL).
The
section
Tulipa
is
found
in
Europe
and
the
Middle
East
(Hoot
1973).
The
species
are
generally
closely
related
to
the
cultivated
tulip,
T.
gesneriana,
T.
rhodopea
is
regarded
to
be
a
subspecies
of
T.
36
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
hungarica,
occurring
in
Rumania
and
Bulgaria,
respectively
(Hoot
1973).
T.
armena
and
T.
galatica
are
treated
as
conspecific,
sympatric
in
Asia
minor,
and
differing
in
flower
colour
(HALL
1940),
chromosome
number
and
amount
of
hetero-
chromatin
(Table
5).
In
T.
galatica
two
similar
and
one
deviating
haploid
set
of
chromosomes
with
respect
to
C-banding
pattern
was
found
(Fig.
11),
which
could
indicate
an
alloploid
origin.
T.
gesneriana
has
crossability
relationships
with
some
species
of
sect.
Eichleres
(VAN
EIJK
&
al.
1991),
but
differences
in
the
mean
amount
of
heterochromatin
occur
between
sect.
Tulipa
and
sect.
Eichleres.
T.
suaveolens
has
been
mentioned
to
be
the
ancestor
of
the
T.
gesneriana
"Duc
van
Tol"
types
(JANsE
1966).
The
large
variability
of
T.
suaveolens
together
with
that
of
the
two
accessions
of
T.
gesneriana
used
will
be
dealt
with
in
a
separate
paper.
In
sect.
Tulipa
a
group
of
Neotulipae
was
described
in
the
nineteenth
century,
mainly
found
on
the
western
and
southern
borders
of
the
Alps,
and
most
of
the
species
known
from
only
one
locality
(LEVIER
1884,
HALL
1940).
Because
of
their
western,
isolated
distribution
they
should
be
regarded
as
garden
escapes
of
T.
gesneriana
and
they
are
treated
as
one
species.
Some
forms
included
in
the
Neotulipae
show
hairy
bulb
tunics
and
red
or
scarlet
flower
colours
with
a
distinctive
black
blotch,
yellow
margined.
These
forms
like
T.
maleolens
should
be
designated
as
part
of
sect.
Tulipanum
(BAKER
1874:
Eriobulbi),
of
which
species
occur
in
the
same
region.
Taxonomic
enumeration
Tulipa
L.
Sp.
Pl.:
305
(1753)
Subgenus
Tulipa.
Basionym:
sect.
Leiostemones
BoIssIER,
Fl.
Orientalis
5,
p.
192
(1882).
Homotypic
synonym:
sect.
Tulipa
MARAIS,
in
DAVIS,
P.
H.,
(Ed.):
Flora
of
Turkey
and
the
East
Aegean
Islands
8,
p.
306
(1984).
Type
species:
T.
gesneriana
L.
(1753).
Section
Clusianae
BAKER,
Gard.
Chr.
n.s.
19:
691
(1883).
Homotypic
synonym:
subsect.
Clusianae
HALL,
The
genus
Tulipa,
p.
80
(1940).
Type
species:
T.
clusia-
na
DC.
(1803)
(lectotype,
designated
here).
T.
clusiana
DC.
in
REDOUTE,
Les
Liliacees
1,
t.
37
(1803).
Heterotypic
syno-
nym:
T.
aitchisonii
A.
D.
HALL,
J.
Bot.
76:
313
(1938;
"aitchesonii").
Forma
clusiana.
Heterotypic
synonyms:
T.
chitralensis
auct.,
Kew
Bull.
app.
2:
53
(1898);
HALL,
The
genus
Tulipa,
p.
85
(1940);
T.
aitchisonii
HALL
var.
clusia-
noides
WENDELBO,
Symb.
Afghan.
4:
162
(1968).
The
flowers
show
a
white
basic
color
with
a
crimson
or
purple
dorsal
band,
and
a
dark
purple
blotch
with
fila-
ments
and
anthers
in
the
same
colour.
The
anthers
may
be
either
shorter
or
longer
(var.
clusianoides)
than
the
filaments.
Forma
stellata
(HooK.)
DASGUPTA
&
DEB,
Candollea
40:
165
(1985).
Basionym:
T.
stellata
HOOKER,
Curt.
Bot.
Mag.
54:
2762
(1827).
Homotypic
synonym:
T.
clu-
siana
var.
stellata
(Hoox.)
REGEL,
Acta
Hort.
Petrop.
1:
54
(1873).
Flowers
possess
a
white
basic
colour
with
a
crimson
or
purple
dorsal
band.
Blotch,
filaments
and
anthers
are
yellow.
Forma
diniae
VAN
RAAMSDONK,
forma
nova.
Heterotypic
synonyms:
T.
fernan-
dezii
BLATT.,
J.
Bombay
Nat.
Hist.
Soc.
37:
420
(1934),
nomen
dubium;
T
porphy-
0.15
_
0.10
Clusianae
Q
Eichleres
Tulipanum
Tulipa
Kolpakowskianae
Tulipanum
0.05
Eichleres
Clusianae
-0.05
Kolpakowskianae
-0.10
Tulipa
Tulipa
subg.
Tulipa
37
-0.15
-0.10
-0.05
0.05
0.10
Fig.
10.
Regions
of
species
groups
in
a
plot
with
the
first
and
second
canonical
variate
based
on
the
combined
dataset
of
1988
and
1989.
1
T.
lanata,
T.
ingens;
2
T.
kuschkensis;
3
T.
eichleri,
T.
micheliana,
T.
fosteriana,
T.
tubergeniana,
T.
hoogiana;
4
T.
greigii,
T.
mogoltavica;
5
T.
vvedenskyi,
T.
albertii,
T.
butkovii;
6
T
sosnovskyi;
7
T.
praestans;
8
T
subpraestans;
9
T.
kaufmanniana,
T.
tschimganica,
T.
anadroma,
T.
dubia;
A
T.
armena,
T.
suaveolens,
T.
gesneriana;
B
T.
hungarica,
T.
rhodopea;
C
T.
marjoletti,
T.
planifolia,
T.
didieri,
T.
grengiolensis;
D
T.
clusiana;
E
T.
aitchisonii;
F
T.
linifolia,
T.
batalinii,
T.
maxi-
mowiczii;
G
T.
montana;
H
T.
oculus-solis;
I
T.
stapfii,
T.
systola,
T.
ulophylla;
J
T.
Julia;
K
T.
aleppensis;
L
T.
praecox;
M
T.
kolpakowskiana,
T.
iliensis,
T.
ferganica;
N
T.
zenai-
dae;
0
T.
ostrowskiana,
T.
tetraphylla,
T.
behmiana;
X
hybrid
between
T.
stapfii
(I)
and
T.
praecox
(L)
reochrysantha
BLATT.,
J.
Bombay
Nat.
Hist.
Soc.
37:
421
(1934),
nomen
dubium;
T.
stellata
chrysantha
HORT
ex
HALL
(1940),
non
BOISSIER;
T.
clusiana
var.
chrysan-
tha
(HALL)
SEALY,
Curtis
Bot.
Mag.
n.s.
16S:
13
(1948),
nomen
nudum.
Tepala
lutea
intus
basi
macula
pollide
violacea
vel
sine
macula
extus
stria
coccina
dorsa-
li.
Holotype:
T.
DE
VRIES
1106,
acc.nr
.
68590,
IVT,
Wageningen;
paratype:
T.
DE
VRIES
1115,
acc.nr
.
70639,
IVT,
Wageningen
(WAG).
Flowers
show
a
yellow
basic
colour
without
or
with
a
light
purple
blotch,
but
with
a
crimson
dorsal
band
on
the
tepals,
and
yellow
filaments
and
anthers.
The
anthers
may
be
either
shorter
(T.
porphyreochrysantha)
or
longer
than
the
filaments.
Cultivar
'Cynthia'
is
included
in
this
forma.
Forma
cashmeriana
(HALL)
VAN
RAAMSDONK,
comb.
&
stat.
nov.
Basionym:
T.
aitchisonii
subsp.
cashmeriana
HALL,
J.
Bot.
76:
313
(1938).
The
flowers
are
com-
pletely
yellow,
sometimes
with
a
brown
blotch,
but
without
a
crimson
dorsal
band.
T.
montana
LINDLEY,
Bot.
Reg.,
t.
1106
(1827).
Synonym:
T
wilsoniana
HooG,
Gard.
Chron.
n.s.
32:
50
(1902).
38
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
40014
Ek
t
i
l
e
V,
Ilea. IWO
a
e
d
ip
I
4111
141
11)
lat
Fig.
11.
Idiogram
of
the
triploid
species
T.
galatica
(84266)
T
linifolia
REGEL,
Acta
Hort.
Petrop.
8:
648
(1884).
Synonym:
T.
maximowiczii
REGEL,
Gartenflora
38:
505
(1889).
Forma
linifolia.
Forma
chrysantha
(BOISSIER)
VAN
RAAMSDONK,
comb.
nova.
Basionym:
T.
chrysantha
BoIssIER
in
KOTSCHY,
Fl.
Pers.
Bor,
p.
78
(1846);
BOISSIER,
FL
Oriental-
is
5:
193
(1882).
Heterotypic
synonym:
T.
batalinii
REGEL,
Gartenflora
38:
506
(1889).
This
forma
differs
from
f.
linifolia
in
having
yellow
flowers
instead
of
red.
Both
colour
formas
are
indistinguishable concerning
other
characters.
T
chrysan-
tha
BOISSIER
was
considered
by
HALL
(1940)
to
be
the
yellow
forma
of
T.
montana,
but
almost
all
accessions
in
our
collection
identified
as
yellow
T.
montana
turned
out
to
be
yellow
T.
linifolia
(Fig.
3).
Section
Kolpakowskianae
(HALL)
VAN
RAAMSDONK,
stat.
nov.
Basionym:
subsect.
Kolpakowskianae
HALL,
The
genus
Tulipa,
p.
134
(1940).
Type
species:
T.
kolpa-
kowskiana
REGEL
(1877)
(lectotype,
designated
here;
=
T.
altaica).
T.
altaica
PALL.
ex
SPRENGEL,
Syst.
2,
p.
63
(1825).
Heterotypic
synonyms:
T.
kolpakowskiana
REGEL,
Acta
Hort.
Petrop.
8:
650
(1877),
T.
iliensis
REGEL,
Garten-
Tulipa
subg.
Tulipa
39
flora
28:
162
(1879),
T.
anisophylla
VVEDENSKY,
Bjul.
Sredn.
Gos.
Univ.
21
(16):
147
(1935).
Var.
altaica.
Var.
ferganica
(VVED.)
VAN
RAAMSDONK,
comb.
nova.
Basionym:
T
ferganica
VVED.,
Bjul.
Sredn.
Gos.
Univ.
21
(16):
148
(1935).
Var.
ferganica
show
wider
stems
and
leaves,
and
larger
floral
parts
than
var.
altaica
(Fig.
4),
and
differ
in
area
of
distribution:
var.
altaica
is
found
in
the
Tien-Shan
mountain
range,
and
in
some
parts
of
western
China,
whereas
var.
ferganica
is
endemic
in
the
Pamir-Alai
mountain
range
and
Fergana
basin
(How
1973).
T
lehmanniana
MERCKLIN,
Mem.
Acad.
Pet.
Say.
Etr.
7:
513
(1854).
Heterotypic
synonym:
T
zenaidae
VVEDENSKY,
Bjul.
Sredn.
Gos.
Univ.
21
(16):
150
(1935).
T.
tetraphylla
REGEL,
Acta
Hort.
Petrop.
3:
296
(1875).
Heterotypic
synonym:
T.
behmiana
REGEL,
Acta
Hort.
Petrop.
6:
505
(1880).
Subsp.
tetraphylla.
Subsp.
ostrowskiana
(REGEL)
VAN
RAAMSDONK
comb.
nova.
Basionym:
T
ostrowskiana
REGEL,
Acta
Hort.
Petrop.
8:
649
(1884).
Subsp.
ostrowskiana
has
more
erect
and
less
undulated
leaves
than
subsp.
tetraphylla
(Fig.
4),
and
differs
in
area
of
distribution:
subsp.
tetraphylla
is
found
in
the
NE
part
of
the
Tien-Shan
mountain
range
and
some
parts
of
western
China,
whereas
subsp.
ostrowskiana
is
found
in
the
eastern
part
of
Turkestan
(Hoot
1973).
Section
Tulipanum
DE
REBOUL,
Giorn.
Bot.
Ital.
2:
60
(1847).
Homotypic
syno-
nyms:
sect.
Eriobulbi
BAKER,
J.
Linn.
Soc.
14:
276
(1874),
subsect.
Oculus-solis
HALL,
The
genus
Tulipa,
p.
104
(1940).
Type
species:
T.
oculus-solis
ST.
AMANS
(1804)
(lectotype,
designated
here).
Series
Tulipanum.
Type
species:
T.
agenensis
DC.
(1804).
T.
agenensis
DC.
in
REDOUTE:
Les
Liliacees
1,
p.
60
(1804).
Heterotypic
syno-
nyms:
T.
oculus-solis
ST.
AMANS,
Rec.
Soc.
Agri.
Agen.
1:
75
(1804),
nomen
nudum;
T.
maleolens
DE
REBOUL,
Nonnul.
sp.
tulip.
not.
App.:
1
(1823).
T.
systola
STAPF,
Denkschr.
Akad.
Wien
50:
17
(1885).
Heterotypic
synonyms:
T.
stapfii
TURRILL,
Curtis
Bot.
Mag.
157:
t.
9356
(1934),
T.
ulophylla
WENDELBO,
Nytt.
Mag.
Bot.
14:
99
(1967).
T.
kuschkensis
B.
FEDTSCHENKO,
Bot.
Jahrb.
50:
612
(1914,
nomen;
1932,
descr.)
T.
julia
C.
Kocti,
Linnaea
22:
225
(1849).
T.
aleppensis
BOISSIER
ex
REGEL,
Acta
Hort.
Petrop.
2:
450
(1873).
Series
Aureo-fasciatae
VAN
RAAMSDONK,
ser.
nova.
Plantae
magnae
caule
longo
et
crasso
foliis
latis
floribus
magnis
et
tepalis
distincte
luteo-fasciatis.
Type
species:
T.
praecox
TENORE
(1811).
The
species
of
this
series
has
a
large
habit
(stem
length
40-55
cm),
thick
stem
(6-8
mm),
broad
leaves
(50-75
mm),
large
flowers
(7-9
cm)
and
a
distinct
yellow
dorsal
band
on
the
tepals.
The
yellow
band
is
very
occasionally
found
in
T
systola.
T.
aleppensis
is
intermediate
by
having
broad
leaves
and
being
triploid
like
T.
praecox.
T.
praecox
TENORE,
Flora
Napolitana
1,
p.
179
(1811).
Section
Eichleres
(HALL)
VAN
RAAMSDONK,
stat.
nov.
Basionym:
subsect.
Eichleres
HALL,
The
genus
Tulipa,
p.
122
(1940).
Type
species:
T.
eichleri
REGEL
(1874)
(lectotype,
designated
here).
40
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
Series
Lanatae
VAN
RAAMSDONK,
ser.
nova.
Tunica
parte
superiore
lanata
indumento
apice
bulbi
visibili.
Basi
tepalorum
macula
nigra
acute
marginata
saepe
margine
lutea
distincta
et
stigmate
perpallide
viride.
Type
species:
T.
lanata
REGEL
(1884).
The
species
of
this
series
are
distinct
by
having
a
woolly
layer
of
hairs
in
the
upper
part
of
the
tunics,
a
tuft
of
hairs
visible
on
top
of
the
bulb,
a
sharply
edged,
black
heart
("blotch")
with
or
without
a
distinct
yellow
border
at
the
base
of
the
tepals
and
a
colourless
stigma.
Only
the
slight
hairiness
in
the
lower
part
of
the
bulb
dif-
fers
from
that
of
the
species
of
sect.
Tulipanum.
The
species
are
related
to
the
next
series.
T.
lanata
REGEL,
Acta
Hort.
Petrop.
8:
647
(1884).
T.
ingens
HooG,
Gard.
Chron.
n.s.
32:
14
(1902).
Series
Eichleres.
Type
species:
T.
eichleri
REGEL
(1874).
Species
of
this
series
show
a
sharply
edged, black
blotch
usually with
a
distinct
yellow
border
in
the
center
of
the
flower,
and
either
a
red-coloured
stigma
(T.
tubergeniana)
or
a
pale
yellowish-green
stigma.
T.
eichleri
REGEL,
Gartenflora
23:
193
(1874).
Var.
eichleri.
Var.
micheliana
(Hoot)
VAN
RAAMSDONK,
comb.
nova.
Basionym:
T.
micheliana
HooG,
Gard.
Chron.
n.s.
32:
350
(1902).
Var.
eichleri
and
var.
micheliana
are
com-
parable
in
morphology
(Fig.
6)
but
var.
micheliana
shows
a
much
dwarfer
habit.
The
varieties
differ
in
area
of
distribution:
var.
eichleri
is
endemic
in
S
Caucasus
and
N
Iran,
while
var.
micheliana
occurs
in
NE
Iran,
extending
to
Pamir
Alai;
a
transient
zone
with
both
varieties
may
occur
in
N
Iran
(HooG
1973).
T.
tubergeniana
HooG,
Gard.
Chron.
n.s.
35:
358
(1904).
Heterotypic
synonym:
T.
hoogiana
B.
FEDTSCHENKO,
Gard.
Chron.
n.s.
48:
53
(1910).
T.
fosteriana
HooG
ex
W.
IRVING,
Gard.
Chron.
n.s.
39:
322
(1906).
Series
Vinistriatae
VAN
RAAMSDONK,
ser.
nova.
Folia
supra
striis
vinaceis
suffulta.
Tepala
macula
fusca
vet
nigra
colore
vivide
et
lucente
rubro
transiente.
Type
spe-
cies:
T.
greigii
REGEL
(1874).
The
species
of
this
series
is
distinct
by
having
anthocyanin
stripes
on
the
upper
surface
of
the
leaves
and
a
brown
or
black
blotch,
fading
in
the
bright,
shiny
red
colour
of
the
flower.
Anthocyanin
stripes
are
only
occasionally
found
in
some
other
species,
like
T.
eichleri
(ser.
Eichleres),
T.
alber-
tii
(ser.
Undulatae)
and
T.
kaufmanniana
(ser.
Spiranthera).
T.
greigii
REGEL,
Gartenflora
22:
290
(1873).
Heterotypic
synonym:
T.
mogol-
tavica
M.
POPOV
&
VVEDENSKY,
Herb.
Fl.
As.
Med.
no.
594
(1935).
Series
Undulatae
VAN
RAAMSDONK,
ser.
nova.
Folia
margine
saepe
undulato.
Caules
breves.
Macula lutea
plerumque
gradatim
marginem
versus
atrans
et
colore
rubro
tepalorum
transiens.
Type
species:
T.
albertii
REGEL
(1877).
The
species
in
this
series
shows
leaves
with
often
undulated
margins,
short
stems
(10-35
cm)
and
usually
a
yellow
blotch,
in
some
populations
gradually
darkening
towards
the
blotch
margin,
fading
away
in
the
red
flower
colour.
T.
albertii
REGEL,
Acta
Hort.
Petrop.
5:
264
(1877).
Heterotypic
synonyms:
T
vvedenskyi
BOCHANTZEVA,
Not.
Syst.
Herb.
Uzb.
14:
3
(1954),
T.
butkovii
BOCHANT-
ZEVA,
Not.
Syst.
Herb.
Uzb.
16:
8
(1961).
Tulipa
subg.
Tulipa
41
Series
Luteo-apiculatae
VAN
RAAMSDONK,
ser.
nova.
Caules
flaccidis
irregulariter
flectantes.
Flores
rubri
macula
nigra.
Filamenta
antherae
pollenque
atropurpurea
vel
nigra.
Type
species:
T.
sosnovskyi
AKHVERDOV
&
MIRZOJEVA
(1950).
The
species
of
this
series
shows
flaccid
stems,
bending
irregularly,
and
red
flowers
with
a
black
blotch
and
very
dark
filaments,
anthers
and
pollen.
The
filaments
have
a
dis-
tinct
yellow
tip.
T.
sosnovskyi
AKHVERDOV
&
MIRZOJEVA,
Tr.
Bot.
Inst.
Ac.
Sc.
Arm.
7:
31-32
(1950).
Series
Multiflorae
VAN
RAAMSDONK,
ser.
nova.
Tunica
bulba
dura.
Flores
rubri
vel
armeniaci
since
macula.
Filamenta
colore
tepalis
simila
rariter
atropurpurea.
Antherae
atropurpureae
vel
luteolae.
Plant
plerumque
plusquam
florem
unum
et
etiam
usque
flores
septem
gerens.
Type
species:
T.
praestans
HooG
(1903).
The
species
of
this
series
is
distinct
by
showing
hard
bulb
tunics.
Flowers
are
entirely
red
or
golden
yellow,
which
means
the
absence
of
a
blotch
and
usually
filaments
with
the
same
colour
as
the
tepals
and
with
dark
purple
or
yellow
anthers.
The
plants
have
usually
more
than
one
and
up
to
seven
flowers
per
stem.
This
charac-
teristic
is
extremely
rare
in
some
other
species
of
subg.
Tulipa,
i.e.,
T.
fosteriana
(ser.
Eichleres)
and
T
kaufmanniana
(ser.
Spiranthera)
may
occasionally
show
two
flowers.
It
was
not
found
in
T.
sosnovskyi
(ser.
Luteo-apiculatae)
as
reported
by
BOCHANTZEVA
(1982).
One
population
in
our
collection
shows
dark
purple
fila-
ments.
Besides
the
type
species
T.
praestans
with
red
flowers
a
second
taxon
endemic
to
Afghanistan
has
to
be
described
possessing
yellow
or
ochre
flowers.
T.
praestans
HooG,
Gard.
Chron.
n.s.
33:
364
(1903).
Series
Spiranthera
(VVED.)
VAN
RAAMSDONK,
stat.
nova.
Basionym:
sect.
Spiran-
thera
VVEDENSKY
(1935).
Type
species:
T.
kaufmanniana
REGEL
(1877).
T.
kaufmanniana
REGEL,
Gartenflora
26:
194-195
(1877).
Heterotypic
syno-
nym:
T.
montana
var.
amblyophylla
POST,
Pl.
Post.
1,
p.
13
(1890).
T
tschimganica
BOCHANTZEVA,
Not.
Syst.
Herb.
Uzb.
16:
10
(1961).
Heterotypic
synonym:
T.
anadroma
BOCHANTZEVA,
Not.
Syst.
Herb.
Uzb.
16:
7
(1961).
T.
dubia
VVEDENSKY,
Bjull.
Sredn.
Gos.
Univ.
21
(16):
148
(1935).
Series
Glabrae
VAN
RAAMSDONK,
ser.
nova.
Facies
interior
tunicae
bulbi
glaberri-
ma.
Etiam
coronula
pilifera
basi
bulbi
absens.
Type
species:
T.
subpraestans
VVE-
DENSKY
(1935).
The
inner
side
of
the
bulb
tunics
of
the
species
in
this
series
is
completely
gla-
brous.
Especially
the
hairy
coronet
at
the
base
of
the
bulb
is
lacking,
which
is
not
found
in
any
other
species.
Flowering
time
is
approximately
one
week
later
than
the
other
species
of
this
section.
T
subpraestans
VVEDENSKY,
Herb.
Fl.
As.
Med.
no
597
(1935).
Section
Tulipa.
Basionym:
sect.
Gesnerianae
BAKER,
J.
Linn.
Soc.
14:
276
(1874).
Homotypic
synonyms:
sect.
Dulipanum
DE
REBOUL,
Giorn.
Bot.
Ital.
2:
60
(1847),
subsect.
Gesnerianae
HALL,
The
genus
Tulipa,
p.
92
(1940).
Type
species:
T
ges-
neriana
L.
(1753).
T.
armena
Boiss.,
Diagn.
2
(4):
99
(1859).
Forma
armena.
Forma
galatica
(FREYN)
VAN
RAAMDONK,
comb.
nova.
Basionym:
T.
galatica
42
L.
W.
D.
VAN
RAAMSDONK
&
T.
DE
VRIES:
FREYN,
Bull.
Herb.
Boiss.
4:
186
(1896).
This
forma
differs
from
f.
armena
in
hav-
ing
yellow
flowers
instead
of
red
ones
with
a
black
blotch.
F.
galatica
is
triploid
(Fig.
11).
Both
colour
formas
are
indistinguishable
as
for other
characters.
T.
hungarica
BORBAS,
Foldmuv.
Erdek.,
p.
561
(1882).
Heterotypic
synonyms:
T.
orientalis
LEVIER,
Bull.
Soc.
Sci.
Nat.
Neuchatel
14:
201-312
(1884);
T.
urumof-
fii
HAYEK,
Verh.
Zool.-Bot.
Ges.
Wien
61:
110
(1911).
Subsp.
hungarica.
Subsp.
rhodopea
(VELENOVSKY)
VAN
RAAMSDONK,
comb.
nova.
Basionym:
T.
rhodopea
VELENOVSKY,
Reliquiae
Mrkvickanae,
p.
28
(1922),
T.
orientalis
var.
rho-
dopea
VELENOVSKY,
Sitz.
Bohm.
Ges.
Wiss.
40:
8
(1899).
Subsp.
hungarica
has
yellow,
or
red
flowers
with
a
black
blotch,
and
in
combination
with
both
flower
colours
yellowish-green
pollen
grains.
Flowers
of
subsp.
rhodopea
are
always
purple-red
with
a
black
blotch
and purple
pollen
grains.
The
subspecies
are
alike
concerning
other
characteristics
(Fig.
9).
The
areas
of
distribution are
different:
subsp.
rhodopea
is
found
in
the
Rhodope
mountains
(Bulgaria),
whereas
subspe-
cies
hungarica
occurs
in
Rumania
and
presumably
also
in
Hungary.
Both
distribu-
tion
ranges
are
divided
by
the
Danube
river
(HooG
1973).
T.
suaveolens
ROTH,
Ann.
Bot.
10;
Neue
Ann.
Bot.
4:
44
(1794).
Heterotypic
synonym:
T.
schrenkii
REGEL,
Acta
Hort.
Petrop.
2:
452
(1873).
T.
didieri
JORD.,
Fragm.
1,
p.
36
(1846).
Heterotypic
synonyms:
T.
planifolia
JORDAN
in
JORD.
&
FOURR.,
IC011eS
1,
p.
19
(1858);
T.
marjoletti
PERR.
&
SONG.,
Bull.
Herb.
Boiss.
ser.
1,
2:
425
(1894);
T.
grengiolensis
THOMMEN,
Bull.
Mur.
Soc.
Val.
Sc.
Nat.
63:
67
(1946).
T.
gesneriana
L.,
Sp.
Pl.,
p.
306
(1753).
Subgenus
Eriostemones
(BOISSIER)
VAN
RAAMSDONK,
stat.
nov.
Basionym:
"sec-
tion"
Eriostemones
BOISSIER
Fl.
Orientalis
5:
196
(1882).
Type:
T.
sylvestris
L.,
Sp.
Pl.,
p.
305
(1753).
The
sections
and
species
have
been
treated
in
VAN
RAAMSDONK
&
DE
VRIES
(1992).
We
express
our
sincere
thanks
to
J.
P.
VAN
DIJK,
W.
A.
VAN
DIJK,
L.
M.
VAN
HEUSDEN,
and
P.
KAAGMAN
for
constant care
of
the
living
plant
collection,
to
all
collectors
for
their
sup-
ply
of
accessions,
to
H.
Q.
VAREKAMP
for
his
invaluable
identifications
of
the
material
used,
to
A.
J.
M.
LEEUWENBERG
(Dpt.
of
Plant
Taxonomy,
Wageningen
Agricultural
Univer-
sity),
for
preparing
the
Latin
diagnoses
and
to
J.
P.
VAN
EIJK
and
W.
EIKELBOOM
for
general
support
of
this
study.
References
BAKER,
J.
G.,
1874:
Tulipa
L.
J.
Linn
Soc.
Bot.
14:
275-296.
1883:
The
species
of
Tulipa.
Gard.
Chron.
n.s.
19:
668, 691,
788-789;
20:
11-12,
71,
153,
168-169,
233-234,
266.
BLAKEY,
D.
H.,
VOSA,
C.
G.,
1982:
Heterochromatin
and
chromosome
variation
in
cultivated
species
of
Tulipa
subg.
Leiostemones
(Liliaceae).
Pl.
Syst.
Evol.
139:
163-178.
BLATTER,
E.,
FERNANDEZ,
J.,
1934:
The
flora
of
Waziristan.
J.
Bombay
Nat.
Hist.
Soc.
37:
419-421.
BOCHANTZEVA,
Z.
P.,
1982:
Tulips:
taxonomy,
morphology,
cytology,
phytogeography,
and
physiology.
English
translated
edition
by
H.
Q.
VAREKAMP.
Rotterdam:
Balkema.
BOISSIER,
E.,
1882:
Flora
Orientalis
5,
pp.
191-201.
Geneva.
Tulipa
subg.
Tulipa
43
BROERTJES,
C.,
HARTEN,
A.
M.
VAN,
1988:
Applied
mutation
breeding
for
vegetatively
prop-
agated
crops.
Amsterdam:
Elsevier.
DASGUPTA,
S.,
DEB,
D.
B.,
1985:
Taxonomic
revision
of
the
genus
Tulipa
L.
in
India
and
adjoining
regions.
Candollea
40:
157-172.
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VRIES,
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CPRO-DLO,
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AA
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Accepted
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28,
1994
by
F.
EHRENDORFER