A new species Tulipa heweri related to T. praestans (Liliaceae)


Raamsdonk, L.W.D. van

Nordic Journal of Botany 18(1): 51-55

1998


A new species of tulip originating from northern Afghanistan, T. heweri, is described (subgenus Tulipa, section Eichleres, series Multiflorae). The species is related to T. praestans and differs by having smaller and less densely-ciliate leaves, yellow or ochre flowers, yellow filaments, anthers and stigma, and a green ovary. The 2 species are allopatric, with both reciprocal crosses between T. heweri and T. praestans yielding a moderate seed set.

Nordic
Journal
of
Botany
A
new
species
Tulipa
heweri
related
to
T.
praestans
(Liliaceae)
L.
W.
D.
van
Raamsdonk
Raamsdonk,
L.
W.
D.
van,
1998.
A
new
species
Tulipa
heweri
related
to
T
praestans
(Liliacceae).
Nord.
J.
Bot.
18:
51-55.
Copenhagen
ISSN
0107-055X.
A
new
species
Tulipa
heweri
is
described
(subgenus
Tulipa,
section
Eichleres,
series
Multiflorae).
The
species
is
related
to
T
praestans
and
differs
by
having
smaller
and
less
densely
ciliate
leaves,
yellow
or
ochre
flowers,
yellow
filaments,
anthers
and
stigma,
and
a
green
ovary.
The
two
species
are
allopatric,
T
heweri
being
endemic
to
northern
Afghanistan.
Crosses
between
T
heweri
and
T
praestans
yielded
moderate
seed
set.
L
W
D.
van
Raamsdonk,
Centre
for
Plant
Breeding
and
Reproduction
Research
CPRO-DLO,
P.O.
box
16,
6700
AA
Wageningen,
The
Netherlands.
Introduction
Tulipa
praestans
Hoog
(section
Eichleres
Boiss.,
series
Multiflorae
van
Raamsdonk)
is
the
only
species
in
subgenus
Tulipa
with
usually
more
than
one
flower
in
the
inflorescence.
Though
species
like
T.
fosteriana
and
T.
kaufmanniana
may
show
two
flowers
occasionally,
T.
praestans
will
show
up
to
seven
flowers
(van
Raamsdonk
&
de
Vries
1995).
T.
praestans
belongs
to
a
monotypic
series
and
takes
an
isolated
position
within
the
section.
Crosses
between
this
species
and
other
rep-
resentatives
of
subgenus
Tulipa
resulted
in
only
some
hybrid
plants
with
T.
gesneriana
(section
Tulipa),
T.
eichleri
and
T.
vvedenskyi
(section
Eichleres),
while
crosses
with
five
other
species
of
section
Eichleres
re-
sulted
in
no
seeds
at
all
(van
Raamsdonk
et
al.
1995).
In
1971
an
expedition
was
organised
to
northern
Af-
ghanistan
by
C.
Grey-Wilson
and
T.
F.
Hewer,
where
a
tulip
population
was
sampled
as
GWH
709,
with
plants
showing
one
to
four
yellowish
or
ochre
flowers
per
in-
florescence.
It
was
presented
as
belonging
to
T.
kolpakowskiana
(Hewer
1974),
presumably
for
the
shape
and
color
pattern
of
its
flowers,
although
T.
kolpakowskiana
is
not
indicated
as
native
to
Afghani-
stan
(Hoog
1973).
The
general
habitus
of
collected
liv-
ing
material
(accession
number
71331)
is
comparable
to
that
of
T.
praestans
plants,
although
a
range
of
specific
Accepted
8-5-1997
CO
NORDIC
JOURNAL
OF
BOTANY
Nord.
J.
Bot.
18(
I)
1998
differences
exists.
The
location,
however,
is
separated
from
the
geographical
distribution
of
T.
praestans
(Pamir
Alai
mountain
range
(Hoog
1973))
at
a
distance
of
about
300
miles.
Investigations
have
been
carried
out
to
study
the
rela-
tionship
between
the
newly
collected
material
and
T.
praestans.
In
this
paper
results
will
be
reported
of
mor-
phological
and
crossability
analyses
in
order
to
estab-
lish
the
status
of
the
new
material.
Material
and
methods
Material
of
ten
accessions
was
used
(Tab.
1).
A
morphological
multivariate
analysis
has
been
car-
ried
out
in
which
most
of
the
above
mentioned
acces-
sions
were
included
(van
Raamsdonk
&
de
Vries
1995).
Some
crosses
have
been
made
in
the
framework
of
the
crossability
program
of
CPRO
(formerly
IVT;
van
Raamsdonk
et
al.
1995).
The
crossability
results
were
analysed
by
means
of
the
crossability
coefficient.
This
coefficient
indicates
the
level
of
interfertility
between
zero
(full
interfertility)
and
one
(complete
intersterility).
A
matrix
of
crossability
coefficients
representing
the
levels
of
interfertility
between
a
set
of
species
can
be
used
for
cluster
analysis,
represented
by
a
dendrogram
(van
Raamsdonk
1992, 1995;
van
Raamsdonk
et
al.
51
Tab.
I.
Accession
number,
chromosome
number,
source
and
origin
of
studied
material
of
Tulips.
No.
Chromosome
no.
Source
and
origin
T.
praestans
Hoog
70626
2n=24
M.H.
Hoog,
Haarlem,
Netherlands
76115
B.
Andreasen,
Denmark;
from
Hort.
Bot.
Oxford
T.
heweri
van
Raamsdonk
71331
C.
Grey-Wilson
and
T.
F.
Hewer
GWH
709:
Salang
Pass,
N
of
Kabul,
Afghanistan
72101
Gibbons,
S0438
77266,
80285
panmixis
populations
of
71331
T.
altaica
Pall.
ex
Springer
(incl.
T
kolpakowskiana
Regel)
68596
2n=24,
48
Thoolen,
Overveen,
Netherlands
74226
2n=24
W.
Kooiman,
Enkhuizen,
Netherlands
74228
2n=24
W.
Kooiman,
Enkhuizen,
Netherlands
76333
2n=24
Hort.
Bot.
Dushanbe,
Tadzhikistan:
original
from
W
Pamir
Alai
82105
Hort.
Bot.
Moscow,
Russia:
original
from
E
Kazakhstan
1995).
This
technique
is
used
to
analyse
the
level
of
interfertility
between
T.
praestans
and
the
newly
de-
scribed
species
(section
Eichleres)
in
comparison
to
T.
eichleri,
T.
fosteriana
and
T
ingens
(section
Eichleres),
and
T.
gesneriana
and
T.
didieri
(section
Tulipa).
Results
and
discussion
The
material
of
GWH
709
collected
at
the
Salang
Pass
was
identified
as
Tulipa
kolpakowskiana
(Hewer
1974;
Rechinger
1990;
now
included
in
T.
altaica,
Van
Raamsdonk
&
De
Vries
1995).
The
description
in
Flora
Iranica
(Rechinger
1990)
fits
that
of
T.
kolpakowskiana,
except
for
the
color
of
the
bulb
tunic,
which
is
reddish
brown
in
T
kolpakowskiana.
However,
Rechinger
(1990)
mentioned
only
one
flower
per
stem
for
the
ma-
terial
of
GWH
709,
whereas
Grey-Wilson
and
Hewer
indicated
up
to
four
flowers
under
natural
circumstances
on
the
labels
of
the
two
herbarium
sheets
in
Kew
(Ap-
pendix
).
The
plants
of
accession
71331
are
character-
ized
by
having
nearly
globular
bulbs
with
dark
brown,
almost
glabrous
tunics,
broad,
ciliate
leaves,
usually
more
than
one
flower
per
stem,
and
obovate
tepals,
yel-
low
or
ochre
with
a
central
reddish
spot
or
stripe.
It
can
be
clearly
distinguished
from
T
kolpakowskiana
by
these
features,
as
is
checked
by
using
living
material
(Van
Raamsdonk
&
De
Vries
1995)
and
herbarium
ma-
terial
of
T.
kolpakowskiana
(Appendix
).
Moreover,
the
geographic
area
of
the
general
species
T.
altaica
in-
cludes
the
Tien
Shan
mountain
ranges
(Fergana
basin),
extending
to
the
Kyzyl
Kum
in
the
west
and
western
China
in
the
east
(Hoog
1973).
It
reaches
its
southern
border
at
the
north
slopes
of
the
Pamir
Alai
mountain
range.
This
area
is
north
of
Afghanistan.
It
can
be
con-
cluded
that
T
altaica
is
not
indigenous
to
Afghanistan,
since
GWH
709
is
the
only
collection
mentioned
for
this
species
in
Flora
Iranica
(Rechinger
1990).
Further
herbarium
material
collected
by
Furse
with
the
same
characteristics
as
the
specimens
of
GWH
709
all
origi-
nate
from
the
Salang
Pass.
Tulipa
praestans
is
characterized
by
having
a
green
stem,
with
anthocyan
coloring
at
top,
ciliate
leaves,
5-7
cm
wide,
red
flowers
with
obtuse
or
acute,
ovate
tepals,
a
small
yellowish
blotch
in
the
center
of
the
flower,
red
filaments,
black
anthers
and
ovary,
and
a
red
stigma
with
black
margins
(Hall
1940;
Stork
1984).
Material
of
accession
71331
differs
from
T.
praestans
in
several
characters,
mainly
concerning
the
flower
color
pattern.
Because
of
these
differences
and
of
the
geographically
isolated
position,
a
new
species
will
be
described.
Tulipa
heweri
van
Raamsdonk,
sp.
nov.
Plantae
fibres
duo
usque
cinque
gerens.
Bulbus
late
ovoideus,
tunicae
nigrescenti-brunneae,
intus
glabrae.
Caulis
viridis
basi
rubens.
Folia
breviter
ciliata.
Tepala
lutea
vel
ochracea
macula
vel
stria
dorsali
rubra.
Filamenta
lutea.
Antherae
et
polline
luteolae.
Ovarium
viride.
Stigma
luteum
marginibus
luteis.
Fig.
1.
Fig.
1.
Photograph
of
the
holotype
of
Tulipa
heweri
(WAG).
52
Nord.
J.
Bot.
18(t)
1998
IIERBARIUM
VAIWNSE
(WAG)
W
L
%As
Itaremthmth
q..
"1.1.
J
.
a
a1
151.4
1.111nuor
To(ersi
lotr•9I
Ramiemulr
Del
I
W
n
vaal
Flaarl.11011
Iv
i
smitvi..41.414,
(111..TA.
k.olutwited
truth
nab
the
w
dd
Afri....terr.
&dun
Pr.
N
rJ
LelvI
Mani
heir!.)
k
th.
rem.
rerJJ1.11
u
Imo,.
Iwo*
10.11.
2,44,
.11.1th,
ohs.
(I
.e..
7
1.
11.ef
ruul
!LI
r
err
...le
Areerl)
hewer.
1.5.
Pima.
on
earl
1
,
1
w.1.'
)rA11111
ON
1,0i
,
orilk
I
414.Mel
1,1+.1
or
writ..
Marlyn.
,410.
eereareir
6
11
1,
Irred,
wine.
.,11)01.4
Arr..
wig..
ACv•MalA
71)7i
in
4
l
r
'
su
Par•rfI.
Mew
•Pit..
7103
UNICATS
TYPE!
Nord.
.1.
Bot.
18(
I)
1998
53
Tab.
2.
Results
of
crosses
between
Tulipa
heweri
and
T
praestans.
Abbreviations:
nkr:
number
of
crosses;
nsuc:
number
of
suc-
cessful
crosses;
npod:
number
of
seed
pods
set;
nsd:
number
of
seeds
obtained;
%:
percentage
of
seeds
grown
to
bulbs;
nb/p:
number
of
F
i
bulbs
per
pod.
mother
T.
heweri
father
T.
heweri
nkr
nsuc
npod
nsd
nb/p
nkr
nsuc
npod
nsd
nb/p
T.
praestans
1
1
11
161
87
12.8
1 1
15
1648
44
14.8
Type:
cultivated
from
seeds
collected
in
the
wild,
Af-
ghanistan,
Salang
Pass,
N
of
Kabul,
van
Raamsdonk
7001-7003
(holotype
WAG,
isotypes:
K,
LE).
Plant
height
10-25
cm.,
bulbs
broadly
ovoid,
bulb
tunics
dark
brown,
glabrous
inside,
stem
green,
reddish
at
base,
leaves
10-18
cm.
long
and
2.5-6
cm.
wide,
shortly
ciliate,
flowers
2-5,
tepals
4-6
cm.
long
and
2-3
cm.
wide,
yellow
or
ochre,
with
a
dorsal
red
spot
or
stripe,
filaments
yellow,
anthers
and
pollen
light
yellow,
ovary
green,
stigma
yellow,
stigma
margins
yellow
(Fig.
1).
Differing
from
T.
praestans
by
the
stem
green
at
top,
smaller
leaves,
ciliae
less
dense,
and
coloring
of
tepals,
filaments,
anthers,
ovary
and
stigma.
Differing
from
7'.
kolpakowskiana
by
its
bulb
shape,
dark
brown
glabrous
bulb
tunics,
ciliate
leaves
and
number
of
flowers
per
stem.
Geographical
distribution.
Afghanistan,
Baghlan
and
Parwan
provinces,
higher
slopes
of
the
Salang
Pass
in
the
Hindu
Kush
mountains,
endemic.
Notes.
The
name
is
given
in
order
to
honour
the
pro-
vider
of
the
material
the
late
prof.
T.
F.
Hewer.
The
available
herbarium
specimens
of
T.
heweri
(Appendix
)
show
a
clinal
variation
with
plants
up
to
35
cm
high
and
one
to
four
flowers
per
stem
at
an
altitude
of
6600-7000
ft
and
plants
of
8-15
cm
high
and
only
one
flower
per
stem
at
10000
ft.
Apparently
the
species
is
capable
of
showing
a
larger
and
multiflowered
plant
habitus
at
lower
altitudes.
Therefore,
a
specimen
grown
from
wild
collected
seeds
in
the
CPRO-DLO
tulip
collection
was
chosen
as
type.
Material
grown
in
a
controlled
environ-
ment
allows
a
better
comparison
with
other
specimens
(i.e.
from
T.
praestans
and
T.
altaica)
which
are
grown
under
identical
circumstances.
Hewer
(1974)
reported
on
some
of
the
accessions
collected
during
journeys
organised
in
1969,
1971
and
1973
in
Iran, Iraq,
Pakistan
and
Afghanistan.
Several
of
these
accessions
have
been
assigned
to
T.
banuensis
Grey-Wilson,
which
was
collected
on
the
north
slope
of
the
Hindu
Kush,
East
of
Banu
and
it
has
been
stated
as
a
relative
of
T.
praestans
(Hewer
1974;
Rechinger
1990).
This
species
was
described
by
having
always
one
flower
and
a
range
of
other
features,
such
as
hairiness
and
color
of
the
bulb
tunic,
smaller
leaves
and
a
differ-
ent
flower
color
pattern,
which
distinguishes
it
from
T.
praestans
(Rechinger
1990).
The
relationship
of
T.
banuensis
with
T.
praestans
is
doubtful.
A
relationship
with
T.
kuschkensis
(section
Tulipanum)
or
with
T.
montana
(section
Clusianae)
could
be
considered.
Material
of
accession
71331
has
been
used
in
multivariate
morphological
analysis
(van
Raamsdonk
&
de
Vries
1995).
Flower
coloring
characteristics
have
not
been
included
in
the
list
of
attributes.
No
significant
dif-
ferences
were
then
found
between
T.
heweri
and
T.
praestans
(van
Raamsdonk
&
de
Vries
1995)
which
is
due
to
the
fact
that
the
characters,
which
afterwards
showed
to
be
discriminating,
were
not
included
in
the
dataset.
Both
reciprocal
crosses
between
T.
heweri
and
T.
praestans
resulted
in
moderate
seed
set
(Tab.
2).
The
number
of
raised
F
1
plants
per
harvested
seed
pod
ranges
low
compared
to
the
numbers
obtained
in
a
set
of
selected
cross
combinations
within
section
Eichleres
(van
Raamsdonk
et
al.
1995:
Tab.
1:
from
5.2
to
106.8
hybrid
plants
per
seed
pod).
Calculation
of
the
cross-
ability
distance
based
on
the
results
of
the
crossing
ex-
periments
with
T.
heweri
and
T.
praestans
resulted
in
a
distance
of
D=0.513
on
a
scale
between
zero
and
one.
The
crossability
coefficient
has
been
applied
to
a
full
diallel
of
twelve
species
of
sections
Tulipa
and
Eichleres
(van
Raamsdonk
et
al.
1995).
Crossability
distances
of
a
selection
of
species,
i.e.
7'.
gesneriana,
T.
didieri,
T.
eichleri,
T.
fosteriana,
T
ingens
and
T
praestans
together
with
T.
heweri
are
used
for
single
linkage
clustering.
The
resulted
dendrogram
is
pre-
sented
in
Fig.
2.
The
crossability
distance
between
T.
heweri
and
T.
praestans
is
larger
than
found
within
the
groups
of
T
gesneriana
and
of
T.
eichleri.
The
specific
status
of
T.
heweri
is
justified,
especially
when
taking
into
account
the
allopatric
geographic
distribution
(Hoog
1973;
Hewer
1974).
Since
it
was
impossible
to
raise
any
intersectional
hybrid
between
species
of
sec-
tion
and
section
Kolpakowskianae
(van
Raamsdonk
et
al.
1995),
the
moderate
crossability
of
T.
heweri
with
a
species
of
section
Eichleres
(
i.e.
T.
praestans)
approves
the
exclusion
of
7'.
heweri
from
section
Kolpakowskianae.
54
Nord.
J.
Dot.
18(1)
1998
T.
gesnerlana
T.
dIdert
T.
elchlerl
T.
foster/am
T.
Ingens
T.
praestans
T.
hewer/
0
0.2
0.4
0.6 0.8
Fig.
2.
Dendrogram
after
single
linkage
clustering
of
cross-
ability
data
of
selected
species
of
genus
Tulipa.
Acknowledgements
I
thank
mr
W.
Eikelboom
for
sup-
port
of
this
study
and
dr
A.
J.
M.
Leeuwenberg
(Dept.
of
Plant
Taxonomy,
Agricultural
University
Wage-
ningen)
for
preparing
the
Latin
diagnosis.
The
curators
of
the
following
herbaria
are
greatly
acknowledged
for
providing
material:
G,
G-BOISS,
K,
L,
WAG.
References
Hall,
A.
D.
1940.
The
genus
Tulipa.
-
The
Royal
Horticultural
Society,
London.
Hewer,
T.
F.
1974.
Some
observations
on
Tulips
in
Iran
and
Afghanistan.
In:
Lilies
and
other
Liliaceae.
Royal
Horti-
cultural
Society,
London,
pp.70-74.
Hoog,
M.
H.
1973.
On
the
origin
of
Tulipa.
In:
Lilies
and
other
Liliaceae.
Royal
Horticultural
Society,
London,
pp.
47-64.
Raamsdonk,
L.
W.
D.
van,
1992.
A
crossability
coefficient
for
the
evaluation
of
crossing
experiments.
Nord.
J.
Bot.
12:
177-182.
1995.
The
cytological
and
genetical
mechanisms
of
plant
domestication
exemplified
by
four
crop
models.
Bot.
Rev.
61:
367-399.
,
Vries,
T.
de,
1995.
Species
relationships
and
taxonomy
in
Tulipa
subgenus
Tulipa
L.
Pl.
Syst.
Evol.
195:
13-44.
,
Eijk,
J.
P.
van,
Eikelboom,
W.
1995.
The
analysis
of
crossability
between
species
of
Tulipa
subgenus
Tulipa.
Bot.
J.
Linn.
Soc.
117:
147-158.
Rechinger,
K.-H.
1990.
Tulipa.
In:
Flora
Iranica,
Cont.
No.
165.
Akademische
Druck-
and
Verlagsanstalt,
Graz.
Stork,
A.
1984.
Tulipes
sauvages
et
cultivees.
Conservatoire
et
Jardin
Botaniques,
Serie
documentaire
13.
Geneva.
Appendix
Herbarium
specimens
investigated
T.
heweri:
Afghanistan:
Baghlan
province,
N
slope
of
Salang
pass,
6600-7000
ft,
C.
Grey-Wilson
&
T.
F.
Hewer
709
(K).
Afghanistan:
originally
from
Salang
Pass,
N
of
Kabul,
cultivated
from
seeds
collected
from
accession
71331,
May
1994,
van
Raamsdonk
7004-
7007
(WAG,
WAHO).
Afghanistan:
Baghlan
province,
N
slope
of
Salang
Pass,
8000
ft,
8-6-1964,
P.
Furse
6618
(K).
Afghanistan:
Baghlan
province,
N
slope
of
Salang
Pass,
9000
ft,
May
1971,
P.
Furse
8809
(K).
Af-
ghanistan:
Parwan
province,
S
slope
of
Salang
Pass,
10000
ft,
April
1968,
P.
Furse
7917
(K).
T
praestans:
Tadzhikistan:
district
Baldschuan,
May
1883
(G-BOISS).
Tadzhikistan:
district
Baldschuan,
Kyzyl-Isu,
8000
ft,
May
1883,
Mussa
s.n.
(G-BO1SS,
K).
Tadzhikistan:
Chodzha-Mumin,
1000
msm,
17
April
1976,
A.
P.
Czukavina
&
N.
M.
Amanova
s.n.
(K).
Turkestan:
no
locality,
1904,
Hort.
Tubergen
s.n.
(K).
Uzbekistan:
Samarkand,
1913,
B.
A.
Fedtschenko
1689
(K).
Uzbekistan:
Samarkand,
12-5-1913,
B.
A.
Fedtschenko
1571
(K).
T
praestans,
cultivated
material.
Netherlands:
Botanic
Garden
Delft,
19-4-1923,
s.n.
(L).
Netherlands:
van
Tubergen
nursery,
12
April
1949,
B.
K.
Boom
B121
(cv.
Fusilier;
L).
Netherlands:
van
Tubergen
nursery,
18
April
1950,
B.
K.
Boom
B2513
(cv.
Zwanenburg;
L).
Netherlands:
originally
from
M.
H.
Hoog,
Haarlem,
ac-
cession
70626,
May
1994,
van
Raamsdonk
7008
(WAG).
Netherlands:
originally
from
Botanic
Garden
Breezand,
accession
70653,
May
1994,
van
Raamsdonk
7009
(WAG).
Netherlands:
originally
from
Botanic
Garden
Oxford,
United
Kingdom,
accession
76115,
May
1994,
van
Raamsdonk
7010
(WAG).
Bulbs
re-
ceived
from
Van
Tubergen,
collected
in
Kew,
Alpine
house,
15-4-1947
(K).
Bulbs
received
from
Van
Tubergen,
grown
at
John
Innes
Hort.
Inst.,
1939
(K).
T
kolpakowskiana.
Kazakhstan:
Tien
Shan
Mts.,
S
of
Alma
Ata,
Mt.
Talgar,
6000
ft,
May
1980,
M.
J.
White-
head
s.n.
(K).
Turkestan:
no
locality,
April
1877,
Werry
s.n.
(K).
Kazakhstan:
Tien
Shan
Mts.,
S
of
Alma
Ata,
Mt.
Talgar,
26-4-1930,
M.
Ganitova
s.n.
(G,
K).
Turkestan:
Darwas,
4000
ft,
Mussa
s.n.
(G-BOISS).
Turkestan:
district
Werny,
April
1877
(L).
Turkestan:
district
Werny,
1885,
Kuschakewicz
s.n.
(G-BOISS,
K).
Turkestan:
no
locality,
30-4-1900,
Willmott
s.n.
(K).
T.
kolpakowskiana,
cultivated
material.
Kazakhstan:
Central
Tien
Shan
Mts.,
vicinity
of
Alma
Ata,
Romanowitz,
cult.
in
Kew,
27-4-1934
(K).
Turkestan:
cultivated
in
Kew,
17-3-1882,
Regel
s.n.
(K).
Nether-
lands:
van
Tubergen
nursery,
12
April
1949,
B.
K.
Boom
B116
(L).
Cultivated
at
John
Innes
Hort.
Inst.,
1939,
Dykes
s.n.
(K).
Nord.
J.
Bat.
18(1)
1998
55
SVEN-ERIK
SANDERMANN
OLSEN
BIBLIOGRAPHIA
DISCIPULI
LINNIEI
BIBLIOGRAPHIES
OF
THE
331
PUPILS
OF
LINN/EUS
.
(
ti
••••••cr
.
.
S
'VeW
:
fl
Ar'-'11
*
ifr""41
67
COPENIIAGEN
1997
Nordic
Journal
of
Botany
NOW
AVAILABLE
170
x
240
mm.
458
pp.
With
small
biographies.
Price
400
DKK
(Danish
Crowns)
From
the
author.
Contact:
e-mail:eso@post6.tele.dk
Sandermann
Olsen.
Vejdammen
8,
DK-2840
Holte.
Denmark
©
NORDIC
JOURNAL
OF
BOTANY
56
Nord.
J.
Bot.
18(11
1998