A
Cytotaxonomic
the
in
of
Viola
Gadella
T.W.J.
and
Museum
(.Botanical
study
Netherlands
Herbarium,
19 th,
{received November
Utrecht)
1962)
Abstract
The
1.
2.
chromosome
numbers
species
of 10
of the
Viola in
genus
the
Netherlands
determined.
were
Viola
has
riviniana
chromosome
various
(most often 2n
40).
3.
It was not possible
numbers:
2n
=
35,
40, 45,
47
46,
=
various
chromosome
4.
Despite
the
material
into
find
to
the
a
correlation
numbers
variability
of
in
V.
between
the
external
and
morphology
riviniana.
it
V. riviniana
proved impossible
the Dutch
divide
to
subspecies.
5.
Some
6.
V.
differential
7.
V. calaminaria
is
canina
not
characters
variable
is not
of
in
V.
riviniana
and
cytological respect
related
to
V. lutea but
to
V. reichenbachiana
in
the
the
described.
are
Netherlands.
V. tricolor complex.
Introduction
Up
to
the
present
genus
Viola
many
species
has
crossing
and
have
some
Clausen
Fothergill
40,
45,
plants of
species
a
pertained
in
and
England
was
studied in
and
in
material
hybridization.
47
respectively
In
certain
“supernumerary”
Schmidt
(1961)
40
often
was
reichenbachiana
in Britain
species
and
their
not
in
and
was
Sweden,
Schmidt
examined.
from
yet
found: 2n
were
40),
British
not
whereas
present.
artificially
produced
have the number 2n
17
=
hybrids
30,
two
plants
6
5,
or
7
According
plants.
of Viola riviniana and
plants
the
In
35,
through
been found in continental
the British
Whereas
=
in South
originated
populations
were
1949,
(1954).
hybrid.
plants of putative hybrid origin, but
descriptions.
all
not
(1932)
obtained. The British
“B” chromosomes
extensively
studied South German
inadequate
of the
examined.
part of the
a
by Sch5fer
numbers
(most
=
plants
or
this has
Valentine described
Viola
were
to
(1954)
was
Germany
different chromosome
and
Schofer
produced abundant seed and apparently had
to
cytological
England by Valentine (1941,
South
Germany only the number 2n
small
only
none
Bruun
quite different from those of
1958)
five
46,
morphological,
In the Netherlands
Germany. The Dutch
1956,
England
few
a
information
species.
(1944)
Viola riviniana
1950,
of
(1931, b), studying the cytology of Viola canina in Denmark,
obtained results
in
numbers
experimental
examined.
cytologically
(1961)
however, only
of the
cytology of the
much
as
moreover,
combination of
a
the
chromosome
available.
the available
area
The
out.
known;
now
data is
cases,
Therefore,
entire
carried
been done,
genetical
In
are
into
investigations
numerous
been
produced
types
of
Schofer
gave
only
artificially
hybrids
were
18
GADELLA
J.
found
to
in South
occur
total, had 2n
of difference is
that the
whereas the British
According
riviniana
woods
ones
and
a
of open
type
(Murbeck) Valentine].
from
intermediate habitats.
neither
riviniana
while
the
subspec.
cytology
distinct
[V
grassland
the
number
Viola
able
calaminaria
determine it
to
but
Westphalia,
the
those from
=
26)
multicaulis
2n
with
logically
Dutch
a
of V.
exact count
V.
(
=
and
Living plants
All
regions.
These
at
15
V.
and
violet-iodine
of
plants
This
(1937)
Blankenrode,
identical with
not
suggested
that
closer
the
(2n
=
counted 2n
calaminaria
=
is,
to
the
were
of the
yellow-
our
Viola
tricolor
48 for
the
determined,
was
not
be
obtain
to
species morpho-
chromosome
cytogeography
found
(1960)
number could
compare
to
V. lutea Huds.
Heimans
exact
are
however, subject
ca.
Lej.).
52, but the
to
48), with which they
conclusion
Finally,
cultivated
studied
was
during
numbers
in order
of these
to
of all
obtain
species.
were
Karpechenko,
stained
according
The
to
in
the
of Utrecht.
Herbarium material
Herbarium.
mitoses
embedded in
portraits
in
paraffin,
haematoxylin
chromosome
Abbe Camera
from
adjacent
Cantonspark
University
the Utrecht
made
some
4 years in the
morphologically.
deposited
counts
method.
an
are
collected in the Netherlands and
were
fixed in
were
the aid
lutea
stagnina))
were
chromosome
/i,
plants
(1936)
calaminaria and
were
plants
of the
most
V.
of
zinc-violets
these
satisfactorily
that Griesinger
the Botanical Garden of the State
Baarn,
The
V.
methods
Only living plants
of
blue
been
never
true
objectives of the present study
complete picture
Material
at
of
Netherlands.
V.
to
prob.
=
tricolor.
species (exept
more
the
(1931, c)
Koch
higher number,
the
floras.
established. One of the
an
variability
investigated, it seemed worth-
morphologically
than
Clausen
as
in
Heimans
are
in
classified
dispute,
of
grounds
zinc-violets
complex (2n
var.
riviniana Rchb. subspec.
.
Lej. This had
morphologically
South
various
flowered
a
type
emend.
a
study of these subjects.
interesting problem in the Dutch violets is the chromosome
of
usually
M.
et
V.
of
however, intermediate types
established, despite several attempts. It is
On
N.W.
of
a
undertake
to
Another
was
point
fertile,
population
morphological
in the Netherlands had been
be
subspecies,
nemorosa
There are,
nor
British
to
sterile.
completely
the
(1941),
Rchb.
minor
As
virtually
were
of the
%
30. Another
=
Germany proved
morphologically
two
(Viola riviniana
had 2n
%
for 90
accounting
one,
10
from
hybrids
Valentine
to
consists of
Valentine)
Germany;
40, the remaining
=
or
were
root-tips.
sectioned
the
gentian
drawn
with
Lucida.
Results
Table 1
gives
a
indication of the
summary
origin
of
of the author’s chromosome counts, with
the
material,
the number of
plants,
and
A
CYTOTAXONOMIG
STUDY
OF
VIOLA
TABLE
IN
THE
19
NETHERLANDS
1
1
or
Species
Plant
Viola odorata
L.
herbaium number
52
33
159, 162
L.
54, 282, 286,
li|
all
Origin
of
3
103
Viola hirta
Number plants
Prov. S. Holland:
near
Rockanje
20
Oisterwijk
20
2
Prov.
IN. Ur a bant:
3
Prov.
Limburg;
6
Prov. Limburg;
4
Prov. Limburg: Eyserbos
3
Prov. N. Holland: dunes
near
Epen
near
near Schin
20
Geul
20
Eys
20
op
near
290
9
and
Bakkum
52,
151
156
4
Prov.
2
Prov.
S. Holland:
S.
between
20
Egmond
dunes
Holland:
near
dunes
Rockanje
20
Oost-
20
near
Voorne
Viola
riviniana Rchb.
A.
The
Netherlands
173
6
Prov.
Friesland:
174
12
Prov.
Drenthe:
near
5
near
180
64
129
130,
136
Prov.
Drenthe;
2
Prov.
Overijssel
:
3
Prov. Overijssel
:
6
Prov.
135
3
Prov.
145
2
Prov.
Oude Mirdum
near
Weerdinge
40
Sleen
47
de Lutte
40
near
Oldenzaal
40
near
Denekamp
40
near
Overijssel:
Overijssel:
between
Denekamp
and Punthuizen
Overijssel:
40
near
Lattrop
40
131
1
Prov.
118
3
Prov.
217
3
Prov. Gelderland: Ermelo (Leuvenumse
213
1
Prov.
4
Prov.
12
3
Prov. IN. Holland:
46
4
Prov.
Overijssel
:
near
Gelderland:
Oldenzaal
near
Berg
46
en
Dal
(Nijmegen)
40
bos)
40
Gelderland :
Putten
(Pinetum
Schovenhorst)
110,
113
Utrecht:
Bergen
aan
S.
45
Rhenen
(Grebbebere)
between
Bergen
Zee
40
Holland:
in
Rockanje,
the
40
152, 153, 207
6
Prov. S. Holland: dunes
157
5
Prov.
S.
Holland:
Rockanje
near
dunes
near
1
50
1
Prov.
1
Prov.
Prov.
N.
40
Brabant:
near
Chaam
Limburg: Schweiberger
47
Bos
near
Mechelen
165
op
198
Limburg:
35
Geerendal
near
Schin
Geul
35
1
Prov.
29
1
Prov.
Limburg:
near
41
1
Prov.
Limburg:
near
4
Prov.
Limburg:
near
2
Prov. Limburg: Geerendal
Limburg:
Savelsbos
near
Rijckholt
119
160
op
193
283
1
Prov.
1
Prov.
40
Oost-
Voorne
297
40
and
dunes
150,
40
35
Vijlen
40
Epen
40
Mook
40
near
Schin
Geul
Limburg:
Limburg:
40
Osebos
near
Eyserbos
Gulpen
40
Eys
40
near
20
T.
W.
J.
1
Table
GADELLA
(Continued)
V
1
s
fe .2
v-
U
Ü
°
V}
.a
u
C
(Ö
a
&
0
cn
«3
9
s,
.c
£
C
317, 354, 373,
Limburg:
Prov.
in
slopes
on
bos”
“Schweiberger
near
the
Mechelen
40
379, 381, 382,
384, 385, 387,
390, 391,
397
3
39
Prov.
Bos
Bovenste
Limburg:
near
45
Epen
17
3
Prov.
Limburg; Ravensbos
near
46
Valkenburg
B.
W.
Germany
235
1
Lower
Saxony: Bentheim
243
1
Lower
Saxony:
C.
201, 202, 203,
7
40
Bentheim
47
Belgium
Prov.
Liège:
Ruïne
Reinardstein
near
40
Robertville
204, 284, 285,
286
287, 288,
289
3
Prov.
Luxembourg:
Vamennes
101
3
4
1
Prov.
A.
Viola
reichenbachiana
Jord.
ex
Between Marche
and La Roche
Luxembourg:
The
Prov.
near
en
en
Ardennes
Bérismenille
63
Prov.
37
2
Prov.
Dene-
191, 192, 194,
5
Prov.
Limburg:
Osebos
1
Prov.
Limburg:
Savelsbos
Overijssel:
Lomanskamp
near
20
Ootmarsum
Onderste
Limburg:
Bos
near
20
Epen
195,
40
20
4
5, 58,
40
Netherlands
Overijssel : Austieberg near
kamp
Bor.
near
Gulpen
20
196
197
near
20
Rijckholt
199, 200
355, 372, 375,
2
Prov.
15
Prov.
376, 380, 383,
Limburg:
Limburg:
Eyserbos
slopes
on
Bos
Schweiberger
near
near
in
Eys
the
20
20
Mechelen
386, 388, 389,
392, 393, 394,
395, 396,
397
B.
190
3
Belgium
Prov.
C.
229, 236, 237,
6
Limburg:
W.
Lower
near
Lanaken
20
Bentheim
20
Germany
Saxony:
near
238, 245, 247
240, 241, 244
3
Lower
Saxony:
Burg
Steinfurt
near
20
Bagno
243, 246
Viola rupestris
F.
var.
2
3
205
W.
Viola
(D.C.)
1
293
Prov.
N.
Prov.
S.
L.
208, 209, 210,
211,
212
5
Prov.
3 km North
Holland:
by
Ir.
Holland:
(collected
Beck.
canina
Saxony:
(collected
Schmidt
arenaria
Lower
by
Friesland:
C.
of Bentheim
Bergen
H.
aan
Doing
Oost-Voorne
20
Zee
Craft)
20
20
Sipkes)
Isle
Schiermonnikoog
s
S-g i
0
374, 377, 378,
Ej
■as
*C
0
16
o
be
I*
5
9
»h
u
■g
of
40
A
CYTOTAXONOMIC
STUDY
OF
Table
or
Species
Plant
herbaium number
1
VIOLA
IN
THE
Number plants
2
chroms e
Diploid number
Origin
of
175
21
NETHERLANDS
(Continued)
Drenthe:
Prov.
Valtherzand
near
Emmen
249
1
96
40
Prov. Drenthe:
40
Oosterhessele
near
3
Prov.
Overijssel:
148
1
Prov.
Overijssel:
179
3
Prov.
Overijssel
1
3
Prov.
9
Prov.
6
4
Prov.
8
1
Prov.
1
Prov.
70, 73
6
Prov.
81
3
Prov. N. Holland:
214
3
Prov. S.
Holland:
Goudriaen
271, 262, 269,
4
Prov.
S.
Holland:
near
Oost-Voorne
40
near
Rockanje
40
84, 85,
87
Utrecht:
near
near
:
Marienberg
40
de
40
Gramsbergen
near
Soestdijk
near
between
Utrecht:
Lutte
40
Maarn
and
40
Maarsbergen
N.
dunes
of
Texel
dunes
of
Texel
dunes
of
Texel
Holland:
40
(de Mok)
N.
Holland:
de
(near
181
N.
40
Koog)
Holland:
(near den Hoorn)
N.
Holland:
40
dunes
Bergen
near
Zee
aan
40
40
St. Maartenszee
near
40
40
270
1
Prov.
S. Holland:
108, 109
6
Prov.
N.
132
3
Prov.
Overijssel :
1
Prov.
Gelderland:
265
Viola
palustris
L.
248
83
2
100
3
Brabant:
Rosmalen
40
Denekamp
48
near
near
Ermelo
near
48
Prov. Utrecht: near Maarn
Belgium.
—
Prov.
48
Luxembourg;
near
48
Mochamps.
Viola
a)
tricolor
subspec.
L.
tricolor
172
177
92,
99
116
8
Prov.
Friesland:
1
Prov.
Overijssel;
near
Marienberg
26
3
Prov.
Overijssel:
near
Ommen
26
Prov.
N.
10
Holland:
Rading
b)
subspec.
curtisii
(Forst.) Syme
and
Oude
near
between
Mirdum
Hollandse
arvensis
Murr.
2
Prov. N. Holland:
10
2
Prov.
48
2
Prov.
S.
Holland:
near
Rockanje
26
43
2
Prov.
S.
Holland:
near
Oost-Voorne
26
128
9
Prov.
S.
Holland:
near
Ouddorp
26
176
2
Prov.
Drenthe :
3
Prov.
Overijssel:
near
Ommen
34
133
3
Prov.
Overijssel;
near
Denekamp
34
114
3
Prov.
Utrecht: near
216
10
Prov.
N. Holland:
102
2
Prov. N. Brabant:
186
2
Prov.
N.
Brabant:
105
2
Prov.
N.
Brabant:
168
6
67, 71
93
Viola calaminaria Lej.
26
Hilversum
N.
near
Holland:
Bergen
near
aan
Egmond
Zee
Belgium.
26
a/d
Hoef
Viola
26
26
—
near
Prov.
Emmen
34
Utrecht
near
near
near
near
34
Hilversum
34
Oisterwijk
34
Valkenswaard
Rosmalen
Liège:
La
Calamine
34
34
52
22
T.
Fig.
1A.
1.
Orig.: Eys;
Rchb.
6.
2n
2n
Viola
=
=
Viola
3.
odorata
Viola
40.
riviniana
47. Orig.:
L.
riviniana
2n
Sleen;
8.
2n
=
Rchb.
Orig.: Rhenen;
Rchb.
W.
=
5.
46.
J.
20.
2n
GADELLA
Orig.: Epen;
=
35.
Orig.:
2.
Viola riviniana Rchb.
Orig.:
Valkenburg;
Viola reichenbachiana Jord.
ex
Viola
hirta
Mechelen; 4.
2n
7.
Bor.
=
2n
=
20.
riviniana
45. Orig.: Putten;
Viola
2n
L.
Viola
=
riviniana
20. Orig.:
Rchb.
Epen.
A
Fig.
1B.
CYTOTAXONOMIC
Viola
9.
Orig.: Rockanje;
palustris
L.
Syme 2n
Orig.;
2n
=
=
rupestris
10.
48. Orig.:
26. Orig.:
Hollandse
F.
Viola
STUDY
W.
OF
VIOLA
Schmidt
canina
L.
2n
var.
=
IN
THE
arenaria
40.
(D.G.)
Orig.:
Maarn;
12.
Viola
13.
Viola tricolor
L.
Murr.
34.
14.
Viola
calaminaria Lej:
arvensis
2n
=
tricolor L.
2n
52. Orig.:
=
la
W.
Beckr. 2n
Gramsbergen;
Oost-Voorne;
Rading;
23
NETHERLANDS
subspec.
subspec.
Orig.:
Calamine.
11.
curtisii
tricolor
Emmen
=
20.
Viola
(Forst.)
2n
15.
=
26.
Viola
24
T.
the
GADELLA
J.
numbers of the dried
collecting
some
W.
general remarks with regard
of the
(or the relation between them)
shows the chromosome
odorata
Viola
a)
voucher
the
to
At
specimens.
the end
morphology and the cytology
species
are
given. Fig.
1
and
(A
B)
portraits of the investigated species.
L.
The chromosome number is in accordance with all data
literature: Clausen
Gershoy
Viola
The
(1937),
and
(1961)
Schmidt
b)
Gorczynski
(1928),
Mangh
(1929),
(1927, 1931a),
hirta
results
(1927,
(1929,
obtained
were
Delay
the
Dutch
reported in the
(1961).
province of Limburg
riviniana
morpholo-
no
could
be
found.
and
V.
reichenbachiana
Rchb.
near
in the southern part
of
Jord.
Bor.
ex
As these
in all
species
were
formerly considered
Viola silvatica Fr.,
species
V.
cases
they
treated
are
reichenbachiana has
the
This in accordance with all other data
1931a), Fothergill (1944),
(1927,
Schofer
no.
2
those
originating from the dunes
and those of the chalk-meadows
Viola
d)
c,
as
Schmidt
(1926),
gical differences between the plants
sea-cost
(1954),
(1948), Fothergill (1944),
of cultivation under uniform conditions
years
the
1930),
(1939).
1931a),
Heilborn
(1934),
After 4
Miyaji
Madge
(1926),
Sghofer
L.
literature: Clausen
Gershoy
Heilborn
(1929),
Theron
cytological
same
given in the
(1948), Fothergill (1944),
Delay
8
Schmidt
(1954),
shows
chromosomes
V.
2n
riviniana,
35,
=
no
plant
small
no.
the other hand,
from
35
50:
5
plant
no.
165;
plant
no.
198:
the
plants
6
showed
species,
has
20.
1950). Figure 1A,
(1949,
whereas
equal size,
the
2n
=
in
In
40).
the
three
were
2n
=
2n
=
2n
plants
small
X
were
=
1
X
were
35,
with
of normal
2
counted:
2n
=
ca.
33
counted:
2n
=
37.
counted:
2n
X
=
the numbers
supernumerary
normal
the fact
1
35,
were
all
Limburg,
present.
were
35,
plants,
of
province
found. All these chromosomes
exactly
7
quite
they
=
various chromosome numbers:
often
metaphase-plates
X
or
Notwithstanding
these
was
exactly
X
other hand,
5,
shows that
(1934), Larsen (1954),
of almost
populations
exactly
12
10
showed
(most
metaphase-plates
6
47
Gershoy
metaphase-plates
X
7
1
the literature: Clausen
given in
are
B-chromosomes
or
4
On
47
different
=
varieties of the
two
larger.
45, 46,
the number 2n
(1961),
be
chromosome number 2n
Valentine
chromosomes
are
on
40,
originating
size;
18
that
to
together. Table
or
34.
2n
=
45,
46
B-chromosomes.
and
All
pollengrains.
that there
resemble each other
are
to
many
a
differences between
certain extent in
general
A
CYTOTAXONOMIC
STUDY
OF
VIOLA IN
THE
25
NETHERLANDS
3-6:
Epen;
1-2:
Orig.:
Ootmarsum.
20).
=
(2n
21-25:
Bor.
ex
Jord.
Gulpen;
18-20:
reichnbachin
Eys;
1 -17:
Viola
of
stipulesRijckholt;
7-10:
the
of
shape
Bentheim;
the
in
Vari ton
2.
Fig.
26
T.
W.
J.
GADELLA
Lat rop;
1-2:
40.
=
2n
Origin: samples:
all
Rchb.
in
rivniana number
Viola
of
region
cauline
mid le
the
of
Chroms e
Mook.
25-26:
We rdinge;
stipules 18-24:
the
of
shape
the
in
Oldenza l;
10-17:
Vari tonDen kamp;
3A.
3-9:
Fig.
A
CYTOTAXONOMIC
STUDY
OF
VIOLA
IN
THE
27
NETHERLANDS
in
Mook;
number
27-29:
Origin:
Rchb.
rivnian
Chroms e
Vijlen;
50-52:
Viola
Rhenen;
of
region
46-49: 40.
cauline
=
Zee;
2n
mid le aan samples:
Bergen
the
of
all
stipules 4 -45:
the
of
Rockanje;
shape
the
3 -43:
in
Dal;
Vari ton
en
Berg
3B.
Fig.
30-32:
28
T.
W.
J.
GADELLA
Schin
Valkenburg,
53-5 :
Origin: 67-70:
45;
=
Rchb.
2n
rivniana Epen,
63-6 ;
V.
of
region
40; 47.
=
2n
=
2n
cauline Sleen,
Bérisménile,
mid le 71-75:
the
59-62: 46;
of
stipules
=
40;
2n
=
2n
the
of
Robertvile,
shape
the
in
Vari ton
56-58:
40;
=
2n
3C.
Geul,
Fig.op
A
STUDY
CYTOTAXONOMIC
appearance. As it is almost
which
among
some
studies
phological
of
The
of
V.
but there is
riviniana,
species
often
are
considerable
a
29
NETHERLANDS
garden conditions.
in many
respects.
agree
smaller
The
overlap.
The characters
plants.
under uniform
of both
reichenbachiana
living
on
constant
characters
vegetative
leaves
THE
to
study some flower characters,
value, in dried material, the mor-
based
only
are
VIOLA IN
impossible
diagnostic
described below remained
The
OF
than
of
those
holds
same
V.
for
true
The
the hairiness of the leaves.
stipules are manifestly different.
Fig. 2 shows the variation in the shape of the stipules of V. reichenbachiana,
Fig.
and
habitats
examples
riviniana
In
(A, B, C) of V. riviniana. Stipules of plants originating from
3
different
riviniana the
V.
of insertion of
In
a
The
fringes
fluctuates
generally, but
with
also
2n
pointed,
number
2n
=
20
partly white,
40,
veins
of
restricted
remain
the
to
of
V.
47
dark
of the spur
of
plants
out
riviniana
one
the
broad
46),
plant
(2n =45),
plant
=
plant
180
no.
213
no.
described
and
is
fill
(2n
=
in
The
V.
V.
of seeds
reichenbachiana,
were
collected
fruits
of
V.
at
as
shoots.
from
Sleen
by
and
The
(2n
=
and
45)
plant
petal
agrees
with
no.
V.
become
petal,
and
are
riviniana
diffuse in
petal. The colour
the
occurrence
plant, plant
purple
purple
of the
veihs of
of the lower
correlated
stouter,
number
V. reichenbachiana
centre
not
part
in
a
the
purple, and
Valentine. The
fewer
petal.
with
with
plants
in
243
plant
(2n
=
no.
47)
of
131
no.
39
and
pollengrains is somewhat larger in V. riviniana
reichenbachiana
number
the
47).
The diameter of the
than
15)
no.
(1949),
provided
dark
the distal
are
(2n =46),
17
no.
is,
plants.
petals
to
chromosomes: the spur is white in
supernumerary
(2n
plants
whereas
have
of the
part
purple-spotted
V.
This
adventitious
the
(white, light
reichenbachiana
the central
have
the lower
spur,
distinctly seperate, those of
usually
are
number,
(Fig. 3A,
40
The venation of the lower
partly purple.)
those
=
Valentine
checked in all
petals,
46,
riviniana branch
V.
per
are
clearly different. Figure 4 shows
lower, spurred, petals. All plants with the
purple
45,
that of the British
whereas
V.
species
two
dark
by
out
not
furrowed spurs of different colour
with
fringes
stipules
chromosome
high
be observed in
however,
was,
a
chromosomes
only
have small
35,
=
the
47, have broad stipules.
=
pointed
was
could
The flowers of the
slender,
of
fringed.
are
stipule at the place
length of that fringe.
riviniana
with the number 2n
As
the differences between the
number
V.
with
plants
supernumerary
180). It
no.
180), 2n
(no.
plants
phenomenon
(plant
In
ones
number of
longer. The
are
species.
for all
stipules.
with
plants
This
are
broad
fringes
both
leaves
a
smaller than the
never
typical
most
uppermost
always, broader than in V. reichenbachiana. The plants
typical
not
for there
in
from Sleen
originating
however
not
the
the
case
every
of
the width of
short;
are
fringe is
V. reichenbachiana the
stipule
In
stipules
entire, but those of the middle
often
are
examined.
were
drawn.
were
(table
per
shows
random
2).
fruit
Fig.
in
reichenbachiana and
5.
is
higher in
Fruits
July in
105
of
from
different
V.
V.
riviniana
than
in
cleistogamous flowers
populations,
riviniana.
viz.
86
30
T.
Fig.
4.
Lower petal and
riviniana
Orig.:
Rchb.
(2-9).
1.
spur
2n
=
W.
of
20.
Epen. Spur white; 3. 2n
GADELLA
J.
Viola reichenbachiana Jord.
Orig.:
=
40. Orig.:
Orig.: Oude Mirdum. Spur purple; 5. 2n
6.
2n
=
40.
light purple;
Orig.:
8.
2n
Schin
=
46.
op
ex
Bor.
(1)
Ootmarsum. Spur dark purple;
=
and
Viola
2n
=
40.
Lattrop. Spur white; 4. 2n
=
40.
40. Orig.:
Geul. Spur purple;
7.
Rhenen.
2n
Orig.: Oldenzaal. Spur white;
Spur purple.
2.
=
9.
Spur white-purple;
45. Orig.: Epen. Spur
2n
=
47. Orig.: Sleen.
A
STUDY
CYTOTAXONOMIC
OF
VIOLA
TABLE
and
Species
2n
20-24
p
24-28
p
THE
IN
31
NETHERLANDS
2
28-32
p
32-36
36-40
,u
40-44
44-46
p
origin
V.
reichen-
20
2
20
2
%
48
%
44
%
%
39
%
59
%
30 %
70
%
2
%
bachiana
(Mechelen)
V. reichenbachiana
(Ootmarsum)
V.
reichen-
20
bachiana
(Bentheim)
V.
riviniana
40
1
%
40 %
59
57
%
35 %
78
%
22 %
48
%
36 %
36
%
38 %
26
%
18
%
48%
28
%
%
(Rhenen)
V.
riviniana
40
3 %
3 %
(Weerdinge)
V.
riviniana
40
(Rockanje)
V.
riviniana
40
8 %
_
6 %
(Lattrop)
riviniana
V.
46
(Valkenburg)
V.
47
riviniana
_
_
2
4%
%
(Sleen)
The
relation
pollengrains
each
(In
Fig.
5.
of
between
100
sample
The
the
9 populations
difference
Rchb.
number
pollengrains
in
of
chromosomes
of V. riviniana
the
and
Rchb.
were
number
of
and
V.
and
the
diameter
reichenbachiana Jord.
of
ex
the
Bor.
measured).
seeds
per
Viola reichenbachiana
fruit
Jord.
ex
between
Viola riviniana
Bor.
The germination of the seeds presented some difficulties. The seeds
only germinated outside the greenhouse after a period of frost. This
is
in
accordance with
Hybrids
between the
an
species
On the island of Voorne
was
found. This
observation
a
were not
by
Gershoy,
met
(1928).
with in the Netherlands.
hybrid between V. riviniana
and
V. canina
plant resembled the V. riviniana parent, but had much
32
T.
and
larger vegetative
hybrid turned
out
all
of cultivation,
The
Viola
e)
The
observations
Gershoy
contrast
the
Contrary
in
did
not
In
differ
cytological
(1929,
Clausen
1931a),
(1961).
Bruun
Love
1932),
Fothergill
(1932),
(1956),
for this
of
J.
Moore
(1957),
species only the number
in
Denmark,
observed.
our
Morphologically
somewhat different from the
was
plants
and
of cultivation under uniform conditions
they
the
other
These
country.
plants
were
plants.
the leaf-base
plants
Clausen in
were
These
cordate,
not
cuneate,
was
from Rosmalen and Oosterhesselen.
plants
all
tall
of
years
from
dune
some
W. Beck.
(D.C.)
with
agrees
Schmidt
pollengrains
parts
4
=
and
the observations
slender, but after
days.
and many
the Netherlands.
abnormal
other
arenaria
var.
20
authors:
Love
the material from Rosmalen
collected
years
few
a
(1961), Schofer (1954), Schmidt (1961), but in
found in
to
no
during 3
mostly shrivelled,
literature:
(1954),
several
(1934),
Harvey
was
material
2n
J. Clausen (1931b,
to
40
=
formed
L.
with
Gershoy
Moore and
2n
canina
accordance
In
(1944)
in
Sghofer
(1934),
were
flowers withered within
Schmidt
number
mentioned
Viola
f)
F.
rupestris
number of the
chromosome
40. No seeds
formed.
were
chromosome
The
parts.
=
quite abnormal,
were
pollengrains
GADELLA
J.
cleistogamous
pollengrains
dwarf
floral
be 2n
to
W.
plants
as
in
supposed
are
belong to the variety dunensis W. Beckr. There are, however, also
plants from the margin of the dunes with cordate leaf-base (plant no.
to
and inland
81)
investigations
dune
our
plants with
needed
are
plants
cuneate
in order
there exists
leaf-base
to
ecoclinic
(plant
no.
214). Further
determine whether
variation with
or
not
to
respect
in
the
leaf-base character.
Viola
g)
In accordance
the
Dutch
Fernandes
Jorgensen,
(1956).
been
with
material
(1950),
all
had
cytological
number
(1944),
(Quezel,
=
48:
Gershoy
with
in
Clausen
Europe,
(1931a),
and Bold
(1958),
Love
the number 2n
(1934),
and
=
Love
24 have
1957).
L.
Lej. (all belonging
species
investigations
2n
Westergaard
and
Viola tricolor
of the
the
Fothergill
Sorensen
The chromosomes of V.
naria
other
Only in North Africa plants
found
h)
L.
palustris
to
tricolor
L.,
V. arvensis
Murr., and V. calami-
the section Melanium)
dealt with in
this paper,
which
are
larger than those
belong
to
the
section
Nominium.
The inland
2n
=
as
well
as
the dune form has
26. This is in accordance with all other
the chromosome number
cytological investigations;
A
Fig.
6.
region
CYTOTAXONOMIC
in
Variation
of
Clausen
Gershoy
Viola
the shape
tricolor
(1921,
STUDY
L.
1926,
of
VIOLA IN
the leaves
subspec.
1927,
OF
tricolor
THE
and stipules
(population
of the
middle
Hollandse
1931a), Fothergill (1938,
Morphologically the
two
kept its ascending
(subspec.
cauline
Rading).
1941,
1944),
(1934).
subspecies
are
clear-cut.
During
cultivation under uniform conditions the annual inland form
tricolor)
33
NETHERLANDS
curtisii)
remained
habit, whereas the perennial
prostrate. Besides,
the
4
years
of
(subspec.
dune
plants
length of the spur
34
T.
Fig.
7.
the
Variation in
of
region
Viola
tricolor
of the dune
plants
appendages,
whereas
long
the
as
calyx
Figs.
region
(plant
no.
Viola
Our
The
of
inland
Fothergill
Love
and
leaves and
a
as
the
Viola
plants
zinc
mitted
of
after
(plant
tricolor
curtisii
long
the
as
calyx
the
4
about as
spur,
dune plants are
years
of cultivation.
116)
no.
and
dune
plants
are
ovate-lanceolate,
are
smaller
and
are
same
(1944),
Love
chromosome number
a
was
1922,
Matsunami
of the middle
shown in
Fig.
tricolor.
The
Viola
cauline
in order
8,
stipules
(fig. 8,
and
previously
1926,
1927,
Hagiwara
no.
2);
to
are
region of collection
permit
a
comparison
sometimes of the
the leaves
are
same
often much
tricolor.
calaminaria
mine in
Kondo,
as
(1956).
foliage leaves
were
whereas
lanceolate.
Murr.
stipules
larger than those of
11
leaves
even
as
smaller
a
by several other authors: Clausen (1921,
with the leaves of V.
j)
plants
subspecies
subspecies
arvensis
number 216
shape
fleshy,
more
samples had the
1931a),
(1956),
The
appendages.
The leaves of the
counted
twice
remained)
cauline
middle
(population Ouddorp).
128).
those of the
i)
Syme
show the differences between the leaves of the middle
and 7
cauline
(and
of the
and stipules
(Forst.)
the inland form has
generally smaller and
6
leaves
curtisii
subspec.
was
GADELLA
J.
of the
shape
L.
W.
Lej.
La Calamine (Belgium) on rubble of
typical Violetum calaminariae. Six plants per-
collected
chromosome
near
counts:
2n
=
52.
A
Fig.
CYTOTAXONOMIC
Variation in
8.
region
the
of
Viola calaminaria
with
the Viola
agrees
with
Sometimes
OF
that
has
of
V.
stipules
for
V.
VIOLA
IN
THE
many
Murr.
lutea,
of the
middle
cauline
(population Hilversum).
characters
morphological
tricolor.
are
35
NETHERLANDS
shape of the leaves and stipules
Viola arvensis
tricolor-complex
the
characteristic
STUDY
For
example,
L.
subspec.
palmatipartite
sometimes
they
the
curtisii
(fig.
are
length
9
in
common
of the spur
(Forst.)
no.
1)
Syme.
which
pinnatipartite,
is
which
36
T.
of
region
is said
a
crenate
character
the
not
the
V.
9
leaves
tricolor.
no.
in V.
present
4),
and
of
stipules
la
(population
the
middle
The terminal lobes of the
according
to
cauline
Calamine).
stipules
Fothergill
(1944)
lutea. After 4 years of cultivation under
these leaf characters
of the spur, width
(length
ters
GADELLA
J.
calaminaria Lej.
(Fig.
uniform conditions
of
shape
Viola
typical for
be
to
be
may
in
Variation
9.
Fig.
W.
and
well
as
as
some
other charac-
shape of the calyx appendages)
unchanged.
remained
Conclusions
In
cytological
material
our
respect
with
largely
agrees
foreign
material.
The
for
number 2n
this
Cytologically
our
material. For the
V. riviniana
reports
a
52
for
V.
calaminaria
the first
represents
no
material of V. riviniana is
explanation
not
discuss
can
printer’s
Contrary
to
these
very close
to
count
the British
of the chromosome number 2n
occurrence
be offered.
this number for material
he does
to
=
species.
plants
=
not
met
number 2n
V.
riviniana
regarded
seed
as
=
as
in
originating from Croxdale, Durham,
further.
The
report
is
due
perhaps
error.
Schmidt,
Schofer and
Valentine,
no
hybrids between
V. riviniana and V. reichenbachiana could be found. Plants with 2n
were
35
Although Valentine (1949)
with,
40
in mixed
even
showed
described
hybrids.
all
by
very
does
clearly
the
The
all
the
not occur
in
plants
characters
therefore,
examined had
artificially
=
plants with
typical
Valentine and could,
Moreover,
production, which
populations.
not
a
30
the
of
be
good
produced hybrids.
A
On
the other
detailed
there
the
1)
material
the
THE
of the
from
be
to
the seed
S.
these
Germany is
30 and 2n
failed
40
=
give
to
In the author’s
putative hybrids.
concerning
of the
production
=
that Schdfer
regretted
37
NETHERLANDS
opinion
plants:
different from
genetically
the
material;
with
plants
The absence of
S.
is
possibilities
two
English
2)
It
descriptions
are
IN
hybrids with the numbers 2n
as
abnormal.
not
VIOLA
OF
hand, in South Germany
plants regarded
was
STUDY
CYTOTAXONOMIC
the number
supernumerary
corroborates
Germany
2n
normal seeds is in better
=
40
of
not
are
hybrid origin.
chromosomes in the material
the first
agreeance
The
assumption.
from
of
production
with the second. It would be
highly
desirable to have detailed
descriptions of some of the hybrid populations
in S.
Germany, in order
to
from
England
The
and
Dutch
material
however,
It was,
populations
woods had 3
the
open
There
regarded
as
did any
type
impossible
seems
have
may
minor
Heimans’s
lutea but
to
have
Some
to
a
morphological
combination of the characters
nemorosa
divide
(DC.)
that
opinion
and minor
by Valentine.
the Dutch material into
these
nemorosa
chromosomes, whereas the plants of the
The
chromosomes.
V. calaminaria should
tricolor- complex
the
that
opinion
Heimans concludes
derived
40
from
plants from
and 4 minor characters.
nemorosa
possess
plants
Dutch
plants
with
chromosomes however, showed several minor characters.
the V.
already of
was
than 40
more
always
type
supernumerary
several
on
from V. tricolor L.
var.
could be
V.
be
not
referred
confirmed. Kloos
calaminaria is related
that
V.
to
V.
calaminaria
V.
to
(1924)
tricolor.
could
be
grounds
alpestris Ging. apud DC. [= V. alpestris
Wittr.]
The
subspecies curtisii of V. tricolor resembles V. calaminaria in many
The number of chromosomes and the
respects.
in
favour
Contrary
to
of
treating
V. riviniana
be observed in
V.
no
calaminaria
as
are
ecological requirements
a
separate
intraspecific cytological
V. canina in the Netherlands.
plants (some of which
are
plant
different
after 4 years
even
while other
characters,
showed 4
11
subspecies. Valentine (1941) suggests that plants of the
two
are
from
plants
conditions.
garden
typical for the subspecies
Therefore it
material
variable.
morphologically
the
minor and nemorosa,
and 5 minor
landscape
is
divide
relation between the environment and
nor
the
comparison with
a
riviniana
to
subspecies
nemorosa
no
characters,
V.
under uniform
dune
was
of
possible
not
into the
of cultivation
facilitate
the Netherlands.
hybrids)
with
species.
variation could
Only in Denmark fertile
more
than
40
chromosomes
present.
ACKNOWLEDGEMENTS
These
investigations
were
carried
out
Lange Nieuwstraat 106, Utrecht (director
made
possible by
(Z.W.O.).
The
for
his
help
with
the
with
with
grant
author
Mennega for their
assistance
a
is
of the
much
assistance
the
the
drawings.
Dutch
in
the
Prof.
indebted
to
Mr.
of the
P.
ofthe
and
for
Systematic
A.
Pure
to
and
to
Mr.
Scientific
partly
Research
Mr.
E.
A.
to Dr. K. U. Kramer,
Mr.
T.
Botany,
were
Florschiitz and
material,
English text,
cultivation of the plants,
for
J. Lanjouw)
Organisation
with the collecting
correction
Institute
Dr.
W.
Fromme
Schipper for
for
his
his
help
38
W.
T.
GADELLA
J.
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