Download Cytotaxonomic notes on some Galium species. A - UvA-DARE

Cytotaxonomic
notes on some Galium species.
A
BY
Prof.
(Communicated by
F.
E.
Kliphuis
A.
Stapleu
at
the
of March
meeting
30,
1974)
Introduction
Within
the
the frame work
following species
were
of
general cytotaxonomic studies of various Galium species
included: Galium rotundifolium L., Galium
arenarium
Galium triflorum Mich., Galium uliginosum L., Galium pumilum Murr., Galium
oelandicum
Ehrend., Galium
tricornutum
Combined
The
results
conditions
of these
Huds.
morphological
during
several
investigations
Galium
Hyl.) Ehrend.,
et
verrucosum
and
cytological
uniform
under
(Stern,
Galium
Dandy,
are
and
studies
Galium
on
Galium
L.
purpureum
made
were
spurium L.,
Lois.,
sterneri
cultivated
plants
years.
described
and
discussed
for each
species
separately.
Material
and
methods
The chromosome numbers determined
The
roottips
In
some cases
plants
Results
a
mitosis.
roottips
fixative, embedded in paraffin,
Karpechenko’s
and stained
number of
to
according
Galium
plants from
mentioned under
are
and
are
indicated
Heidenhain’s haema-
rotundifolium
section
by
population
one
the
an
investigated.
was
collection
same
number.
Such
asterisk.
discussions
rotundifolium L.
Galium
This
micron
numbers
collection
1.
15
on
method.
toxylin
These
fixed in
were
at
sectioned
made
were
is
L.
belongs
characterized
whorls of four, and
1).
(fig.
to the section
by
hermaphrodite
plants
Platygalium (DC.) Koch.
three-veined leaves
having
in
flowers with white corollas in terminal
panicles.
Galium
East
a
rotundifolium has
pubescent,
to
to
40
Asia
cm
Flowers
loose
often
up
to
panicle,
Fruits
2
mm
3
Minor into
an
erect
the
Central, South and South-
Caucasus.
four-angled
stem,
It is
a
perennial
glabrous
cm
long
and 0.5-0.8
turning reddish-brown,
mm
in diameter in
greenish white
or
cm
wide, widest
and black
ovate
long, densely covered with
with
just
ovate
above
when dried.
spreading axillary
white,
with
sometimes
or
high. Leaves in whorls of four, short-petioled,
1-1.5
sub-orbicular,
the middle,
a
and
Europe
creeping stock and
its distribution in
cymes
apiculate
uncinate hairs.
forming
lobes.
346
Galium rotundifolium
Fig.
1.
L.
347
Flowers
The
from
collected
from
middle of
different
several
during
the
is uniform in its
species
Cytological
years
of the
parts
in the
investigation
until
May
July.
morphology. This
experimental
this
on
et al.
diploids
do not
regions
Himalayas
to the
belong
Ehrendobfer
and
the
(1973)
closely related species.
area
Origin
of
the
Andorra: K-1019,
Austria;
wald,
Heide,
Ponta
near
K-1267,
10.3)
d’Envalina.
Nieder-Oesterreich; K-1083,
Herschafts-
Dubner
Steinach, Ober-Franken, Bavaria; K-717,
near
do
Serra
Geres,
Geres; K-1268,
Jihlavsk6
:
Wallis; K-1238,
K-714,
K-525,
near
arenarium
Cytological
K-713,
Zagreb;
Lois.
investigations
hexaploids with
(fig.
on
number could not
made
were
collected
(Lesconil:
Mettmen,
ca.
of Moravia;
Eastern
1360 m, Glarus.
K-569,
Posazavi,
part of Bohemia;
K-728,
K-281,
Serajewo.
near
2).
plants of Galium arenarium Lois, showed
chromosomes.
In the literature Faqerlind
The counts
Leonte, Serra do Geres.
2n =66 chromosomes. The chromosome
without additional
tetraploid
Rio
Bohemia.
Predjumavi,
Galium
near
Northern part
Vrehy, Bohemia;
Yugoslavia;
near
K-1269,
Tatri, Dolina Lejowa.
K-696,
Switzerland: K-566,
in
the
on
(Fig.
(1937) reported
be
2n =44. In
material grown up from seeds
K-700;
is
regular
our
study
this
confirmed.
wild in France
K-295, K-561,
portrait
10.1)
of
the
Plougernan:
as
Department
K-554 and
wel
as
from
of Finistere
Cinder:
K-697),
Department of Morbihan (Plouharnel: K-556) and of the Depart-
ment of Loire
and of the
Messagnes
Atlantique (near Nantes: K-312, K-350,
Department
K-1260;
plants
of Les Landes
Vieux-Boucau
Plage:
K-1265 and
The
to
Bad Diiben.
Tsjbchoslovakia
and
belong
to
portrait is regular without satel-
d’Homen, Serra do Geres; K-1047, K-1270,
Poland:
of the
According
these authors
Oberwart, Burgenland.
Portugal:
plants
these
investigated:
K-289, Manhartsberg,
Germany: K-545,
2.
by
reported
However,
rotundifolium.
plants investigated
material
and Hsu (1968)
respectively.
of Galium
(Fig.
also found
was
(1964) in plants from different
Taiwan
The chromosome
additional chromosomes.
or
exception
showed without
parts of Poland. Khoshoo and Bhatia (1963)
from the
plants
on
cultivated
garden.
material
by Piotbowicz (1958) and Skalinska
and
area
with 2n =22 chromosomes. This number 2n =22
diploids
lites
confirmed
was
distributional
K-1231;
les
St.
(Labenne
Bains:
Giron
K-1261,
Plage:
K-474 and
K-522)
Oc6an: K-1230, K-1259
K-1262
K-1264;
and
K-1263;
Mimizan
Plage:
K-1266).
were
cultivated
during several
years
in the Botanical Garden
348
Galium
arenarium
Fig.
2.
Lois
349
of the State
University of Utrecht,
which
K-1266,
constant
a
of
years
Galium arenarium is
France
showed
remarkable
a
cultivation.
10-20
San Sebastian in the North of
stem
high,
cm
of the coastal
typical representative
a
from the Channel to
perennial,
1973). The plants
K-1259-
nrs.
morphology. All characters mentioned below remained
the
over
collected in
were
in their
uniformity
of the
(with exception
to
prostrate
regions of
It is
Spain.
reddish at the
ascending,
base, glabrous, smooth, quadrangular with ribbed angles, much branched,
with
to
leaves
numerous
1.5
wide, widest
just
stout
at the end of the
cymes
Petals
acute
black,
shining.
Flowering
period:
obtained
from
gardens
nature both
with
and
In
and 0.3
up to 4
mm
(fig.
rarely
in short
3
to
mm,
3).
seed
two
from the botanical
received
samples
and
(Canada)
Lund
is
portrait
these
respect
the
triflorum
is
in
given
fig.
chromosomes
accessory
Over
K-353,
(K-722,
chromosomes.
no
cm
in diameter.
fruit
short,
and
(Sweden)
collected
the North-American continent turned out to be
morphological
characters.
Michaux
from
The chromosome
pattern
deep yellow,
mucronate. Pedicels
not
Montreal
on
2n=66
long
cm
juin-september.
Galium triflorum
Plants
rarely
at the base. Intemodes
to 0.8
teeth, fleshy, darkgreen, shiny. Flowers
branches,
obtuse,
to
10,
above the middle, mucronate, linear-lanceolate to
linear, bearing small
3.
apically, usually defoliated
Leaves in whorls of 5-8
cm.
of
years
in
hexaploid
respectively).
10.6. It shows
hexaploid
plants
cultivation
a
normal
regular
detected.
could be
these
similar in
are
characters
their
remained
constant.
Galium
(Northern-Norway,
Switzerland,
and
this
Within
plants
reported
diploids
The
finding
species
of
a
the
distribution of
series
level
cytotypes
certain
as
and
a
Northem-Europe
and
occurs
also in
Sorsa
occurs.
Khoshoo
some
parts of
relic.
glacial
(1963)
and
found in
Bhatia
(1963)
Himalayas.
in
separated parts of the
of the
area
correlation between chromosome number and
chorology. However, should
about the
considered
tetraploid
from
it
Finland),
polyploid
a
of these
indicates
plant from North-America
there it is
species
Finland
a
Sweden and
this be the case, then still too little is known
plants of different ploidy
level
to
give
a
clear
picture.
Galium
ascending
prickles.
triflorum
or
a
perennial,
Leaves in whorls of 6-8,
ovate-lanceolate,
clear
is
erect, quadrangular,
midvein,
margins rough
Flowers
sessile
or
the
30-60
angles
large,
1-3
mostly
high.
Stem
sometimes
cm
decumbent,
rough by
long and
to
1.2
shortly petiolate, mucronate, usually
otherwise reticulate-veined,
by
cm
antrorse
prickles
or
glabrous, mostly
retrorse
cm
wide,
thin with
dark
green,
smooth.
in cymes of three in theleaf axils.
Corolla small,
greenish.
350
trif lorum Michaux
Galium
Fig.
Fruiting pedicel straight,
to
5
mm.
3.
Fruit
densely covered
with
uncinate
hairs.
Flowering
Galium
4.
time:
uliginosum
From each of
lands and
from
transported
July-August.
L.
(fig.
4).
eighteen populations of
one
to the
in the
Belgian
to
be
experimental garden.
tetraploids
with
uliginosum
Ardennes 7-8
in the Netherlands and those from the
out
Galium
2n =44
The
plants
plants
population
chromosomes.
in the Nether-
were
dug
out and
from
two
from
Belgium
The
populations
plants
turned
from
the
351
Galium
uliginosum L.
Fig.
other
sixteen
Plants from
populations
collected in
collected in the wild,
Norway
in the literature.
diploids
with
location in Denmark and
one
tained from seeds
were
4.
were
also
Diploids by
were
from
2n =22
one
diploids.
diploids. Both cytotypes
Homeyer
(1932)
chromosomes.
in France
all
Two other
are
also
ob-
plants
reported
in material from unknown
origin; by Hancock (1942) for Britain; for Central-Europe by Garajova
(1959) and Ehrendorfer,
lands
by
by
Kliphtjis
Love and Love
(1962)
(in
Love and
Love,
1961)
and for the
and Gadella and Kliphuis
(1956) for Iceland, by
Rohweder
Nether-
(1963), tetraploids
(1937) for Germany
352
and from
htjis
the Netherlands
and
(1932)
Fagerlind
(1934,
satellites
or
Galium
could be observed
uliginosum is rather
height of the plants,
leaves, petals and fruits,
values
plants
calculated
SE =0.22
and
X =30.42
distribution
decumbent
in
1
to
be
and
diploid
plant
respectively)
a
of
a
10
It
is
experimental
factor.
discriminating
The
X =30.15
are:
observation:
fi
N =200;
±
and
respectively. (Number
European species
with
10-60
perennial
a
glabrous,
weak,
little variable.
are
plants).
Central-West
Scandinavia.
as:
of the
size
level. This could be
ploidy
tetraploid
number
a
B-chromo-
cm
quadrangular
a
northern
high.
with
Stem
four
ribs
prickles.
whorls
of
long,
to
up
2
veined,
one
to
7
small
cm.
the
glabrous,
Anthers
and
long
cm
Pedicels
axillary
acute.
period:
the
5-8,
up
in
ovate,
mm
of
no
10.8b
several years in the
during
prove
with
ascending,
Flowers
Flowering
Origin
L. is
mucronate,
lobes
Fruit
not
the
Homeyer
morphology. Characters
correlated with
stomata per
Internodes
flowering.
corolla
20
into
or
with retrorse
prickles.
did
for
uliginosum
lanceolate,
its
± SE =0.23 and SD =3.23 with N =200
n
Leaves
not
cultivated
SD =3.10
of observations:
Galium
(figs.
constant in
regular,
are
10.8a and
and the indument of the
are
Stomata size
garden.
portraits
length of the internodes and pedicels,
Differences if present,
demonstrated on
Gadella and Klip-
reported by
were
1937).
The 2n =22 and 2n =44 chromosome
somes
and
by Kliphuis (1962)
(1963). Tetraploids from unknown origin
2.5
margins
0.5-1.5
mm,
white,
panicles,
mm
3
wide, linear-
with recurved
deflexed after
mm
in
diameter
yellow.
glabrous, rugulose.
July-August.
material investigated:
Diploids:
Denmark: K-206,
France:
K-786,
Jutland, Igleso
La
The Netherlands
:
Roche
en
K-247*,
near
Brenil.
near
Bold Skov.
Dept.
Duurswoude,
K-630*, K-631*, K-633*, K-634*, Speuld,
K-660*,
K-666*,
Holland; K-152*,
K-960*,
Oost
near
prov.
“de
Muy”,
Isle
between Maarn and
Eempolder,
near
Eemnes,
of
prov.
of
Gelderland;
Putten, prov. of Gelder-
Tessel,
prov.
prov.
of Utrecht;
of
Noord-
of Utrecht;
K-37I*, K-372*,
K-373*, K-1007*, former airstrip,
K-374*,
“de Beer”,
Goeree,
Zuid-Holland.
K-1148, roadside
near
Hovringen,
near
South of Dovre.
Tetraploids:
Belgium:
of Friesland; K-189*,
near
Oost Voorne, prov. of Zuid-Holland;
Norway:
prov.
Maasbergen,
prov.
Voorne, prov. of Zuid-Holland;
of
d’Or.
Arboretum: “De Schovenhorst”,
K-190*, Putten,
land;
Cote
K-407*, Robbertsville,
Ardennes.
Dombas; K-1156,
2 km
353
The Netherlands;
between Ermelo
5a.
Galium
(Stem,
et
(Stern,
Weerdinge,
Ehrend.
Hyl.)
Hyl.) Ehrend.
of Drenthe;
prov.
prov.
of
are
(figs.
(Jordan,
5a, b,
has tried to
closely
with
clarety
more
regard
distinguished
sponding
often
related
species
overlapping
1915). Sterner (1944)
bring
a
with Galium
to
its
which
forwardly all were
in
in the
taxonomy of
European
Galium oelandicum
Galium pumilum
(Sterner et Hyl.) Ehrend
Murr.
5.
characters.
complex species,
Europe.
morphologically
and
et
morphological study
this
in North-Western
group
pumilum Murray s.str.,
Fig.
complex have
morphological
well founded
a
pumilum has
1909; Briquet, 1900; Briquet
representatives
central
Galium
c).
pumilum Murr.. Confusion around Galium
unclear and
5c.
sterneri Ehrend. and Galium oelandicum
1846; Braun, 1885; Schuster,
Cavxllier,
especially
on
K-188*,
Gelderland.
been great because taxa of various rank within this
been based
He
et
pumilum Murr., Galium
included in Galium
always
near
Nieuw-Milligen,
pumilum Murr.; 5b. Galium sterneri Ehrend. and
oelandicum
Galium
K-32*,
and
a
corre-
Northern-European
Galium sterneri
Ehrend
354
taxa
the
on
subspecific level,
(0. Dahl) Nordh.;
slesvicence
asp.
with
Stern,
Of
the
variety
tetraploids, diploids
is
distinction
in Britain in
occur
the
largest
The
always
The
areas.
On
by
diploids
are
such
is
1957).
for that
cytotypes
sterneri
by
the other hand
1955,
(Goodway,
the two
ssp.
Isles. This
Ehrend.)
degree of overlapping
possible.
Isles
subspecies:
one
most
extensive
investigations
a part of a
pumilum belongs. (1949,
Ehrendorfer,
that Galium
Galium
as
study
reason
are
Tutin and War-
(Clapham,
1953, 1954,
forms
pusillum complex.
through
This
regular
Leptogalium
1958,
1956,
investigations,
of
part
a
complex
expanded from
perfectly
1955,
the
is
a
an
diploid
level
in Galium (Ehrendorfer,
barriers
in
becoming lesser
has been
able,
with
expand
Galium
hexaploid
CentralIt is
through
pumilum
of
species
West
non
all
nearly
partly
autoploidy,
its
occupy
present
Galium
pusillum
flowering,
smooth,
white,
to
Flowering
Plant
to
pumilum
slightly
and
strong barriers
1973). These
1972,
the
effectively
the
means
complex
to
(Ehrendorfer, l.c.).
as
the
octoploid, rarely
it
and
complex,
occurs
in
glabrous
hairy,
or
linear-falcate,
4
mm
in
leaves
to
in
to
erect
whorls
stems.
of
6-9,
oblanceolate-falcate,
Stems
1-2
cm
the upper
mucronate.
diameter. Fruit
with
tiny papillae,
almost
mm.
time:
remaining
Galium
species
1.5
to
large
through alloploidy,
perennial with ascending
long, the lower narrowly oblanceolate
Corolla
area.
the
number
Europe.
caespitose
linear-lanceolate to
such
Murr. should be understood
the
are
1970,
partly
complex,
a
normal phenomenon
a
polyploids,
higher ploidy levels. By
time and
and
a
the
the lower
in
1963).
higher ploidy levels,
There
1955, 1958; Kliphuis,
effective
extremely
are
larger
to
against intercrossing between the various cytotypes,
1962,
the conclusion
to
aggregate of
by
to which Galium
1960,
came
much
series.
euploid
1946 onwards
made from
were
of the section
result of his
a
pumilum s.l.
of taxa that have
basic
comprising
area
sterneri
tetraploids.
variable and the
not
suecicum
mainly represented
disjunct
slender habit than
very
are
Ehrendorfer as
5a.
and asp.
1962).
burg,
often
has
specific rank (as Galium
the Flora of the British
united under
septentrionale Stern.;
Hyl.
Galium sterneri Ehrend. is
a more
tetraploids
a
et
Stem.)
septentrionale
elevated to
was
smaller and of
In
variety (sap. normanii
ssp.
Denmark and the Northern
part of the British
Ehrendorfer (1960).
that
vestrogoticum
morphologically different
one
Hyl.;
oelandicum Stern,
ssp.
these the subspecies
subspecies
of them with
one
islandicum Stern, et
ssp.
Stern.;
South-Norway,
the
and
comprising six geographically isolated
group
May-August.
green
acid
which
when
dried.
Murr. occurs,
soil,
belongs
in
to
on
dry
grassland
the
to
moderately
vegetations
Mesobromium
dry
which
well drained
are
(Ehrendorfer,
rich
in
1953).
355
because of
Probably
is restricted
species
Galium
faithful
pumilum
of
species
hellinggrasland”)
Plants from
chromosome
is
kind
a
belonging
the
and the
regular,
France:
Netherlands
2n =88
idem,
population
7-10
The
plants
were
were
plants
transported
morphological
to the
characters
plants
were
of
investigated:
K-169,
K-170,
Thoisy
K-785,
slope
Galium
Ten
K-171*,
la
near
Berchere
idem
K-992*,
near
Baelartshoven.
(Dyon).
somes.
Ehrend.
taken
from
These plants
(fig.
flowering
(fig.
shoots
three
populations
is
portrait
10.7b).
or
hairy.
Leaves in whorls of 7-8,
linear,
The
Fruits
Origin
pyramidal
to
of
Denmark,
1.25
in
cm
caespitose,
relic
pedicels
Vorgod
(Sterner
mm.
non
erect, up to
long.
cm
wide, oblanceolate
mm
retrorse
1-2
acute
prickles.
Inflorescence
forming
on
near
et
from
seeds)
the coast; K-1199*,
Hyl.)
Ehrend.
(fig.
K-1195*,
near
Logstor.
5c).
(Sweden). It is
a
diploid,
1960).
perennial, densely caespitose,
with
tubercles.
(obtained
the isle of Oland
(Ehrendorfer,
ascending
four
or
investigated:
K-233*,
is endemic
a
perennial,
1.5
chromosomes
accessory
July.
material
Jutland:
The plant is
shoots
a
long, and
long, glabrous with
mm
Galium oelandicum
postglacial
with 2n =44 chromo-
panicle.
June and
the
species
is
Internodes up to
to 1.5
Blavand, Dunes K-1196*, Klim,
This
regular,
no
and five
investigated cyto-
were
tetraploids
margins with curved,
mucronate,
Flowers
K-995*,
of Galium sternen
Denmark
tetraploid
Flowers to 4 mm, creamy white,
compact
K-994*,
prostrate, the flowering shoots ascending
glabrous
cm,
K-993*,
5b).
turned out to be
The chromosome
observed,
province of Limburg, Schwei-
Kunraderberg;
Eyserbos.
sterneri
plants
logically.
5c.
satellites
or
out and
dug
retained their
obtained from seeds all from Jutland,
a
found. The
were
10.7c).
berg; K-991*,
to
investigated
were
chromosome
B-chromosomes
no
The Netherlands: K-990*, South of the
25
krijt-
importance and consisted largely in the indument and the plants size.
Belgium;
were
(“het
as
Koelerio-Gentianetum.
the years of cultivation.
Existing differences among the
Material
5b.
distinctly basic soils.
limestone-grasslandvegetation
to
France
is
experimental garden.
minor
of
or
the
circumstances
by Westhoff and Den Held (1969)
regarded
Belgium,
(fig.
climatological
in the Netherlands to neutral
portrait
From each
during
favourable
Only octoploids with
cytologically.
observed,
less
up to
short intemodes. Leaves in
15
cm
high, flowering
whorls of 6-9, linear-
356
and
1.5
panicles. Pedicels short,
up
oblanceolate,
midal
Two
plants
logically.
1
cm
were
spurium
Seeds from Galium
Botanical Gardens
were
also
(K-488
diploids
L.
(fig.
with
wide. Flowers white,
mm
to
0.5
and
Fruits
K-779)
2n =22
were
in lax pyra-
to
up
1
mm
investigated
chromosomes,
long.
cyto-
10.7a).
(fig.
6).
spurium collected
of Udine
mm.
(Italy),
in nature and
Stockholm,
received from the
Erevan
and
Leningrad
sown.
Counts
these
long
Oland
from
Both
Galium
6.
to
on
metaphaseplates of roottip mitosis from plants obtained from
seeds,
reported
showed
by
the
chromosome
Fagbblind
(1934,
number
1937)
in
Galium spurium L.
Fig.
6.
2n =20.
material
This
number
from
is
unknown
357
origin and by Podlech
the
Andarab
in
valley
and
Dieterle
Baghlan,
Afghanistan.
remarkable in the genus Galium, because the
number in this
is
genus
X—11.
number X =10 is known,
Galium
molluginoides
The chromosome
in the
ences
H.
portrait is
this
Gadblla and Kliphuis,
the
Love and
1949;
ques,
plant
same
with
aparine
which
1963)
Love,
Podlech
1963;
2n =64
chromosomes,
Gadblla and Kliphuis,
Galium
The
spurium
plant is
quadrangular
glabrous
an
and
Central
rough
on
flowered
four
Pedicels
Fruits
hairs
cymes,
1
in
mm
mm
in
Galium
are
present
Rahn,
1955;
with decumbent
6
retrorse
or
prostrate
Nodes
prickles.
cm.
prickles.
Flowers
in
greenish-white
or
mucronate,
axillary few
white with
long, black, rugulose, glabrous
fruits
Gabriels
and
the
there
(1965)
indumente,
subspecies
uncinate fruit-indument.
e.g.
is
spurium
and
Material
fruits,
infestum
According
and
to
to
are
present,
axillary
flowers
Udine,
Val
Sweden: K-455,
Carlshas.
U.S.S.R.:
Esthonia;
K-268,
this
the
with
Armenia,
et
Kit.)
subdivision
subspecies
Galium
Tellina.
K-318,
characterized
subspecies
(Waldst.
investigated:
K-301,
with white uncinate
subspecies spurium
related to Galium
closely
differences
two
are
the
gated plants used in this study belong
morphological
or
tuberculate bases.
non
to
Italy:
even
of
straight.
differences in fruit
flowers
Larsen and
by
diameter,
period: May-September.
Galium
1971),
long, linear-lanceolate,
retrorse
According
with
1967; Gadella and
Strid,
large chromosomes
high,
to
cm
Flowering
by
1967;
(Pou-
lobes.
1.5-3
with
brous
1962,
occurs
metaphaseplates
edges
internodes up
margins rough by
(3-9)
acute
the
is
America.
cm
the
Leaves in whorls of 5-10,
to 3
up
midrib and
1962,
plates (Bocher,
up to 100
annual,
glabrate,
or
2n =64
through the whole of Europe (except the artic
North
stems,
the
two
fusion
(1955)
1963).
occurs
West-Asia,
parts),
In
differ-
Bocher et al.
the number 2n =66
and
1969).
10.4). No striking
1969;
is
the basic
Hausskn.
1955; Kliphuis,
Dieterle,
1967).
species
et
chromosome number
1956; Kliphuis,
(Kliphuis,
2n =20
chromosome
present and, therefore,
are
but also
and
absent in the 2n =66
are
the
species
basic
and Dieterle,
(Podlech
regular (fig.
very
Larsen and Rahn,
(Bocher,
common
Kliphuis,
L. Krause
spurium from
number
other
two
suggested for Galium aparine by
as
In
unlikely.
seems
Only from
The
common
e.g. Galium ibicinum Boiss.
length of the chromosomes
of chromosomes
most
E.
in Galium
(1969)
aparine.
spurium
2-3
with
K.
gla-
Janchen
the investi-
infestum.
However,
having
leafy bracts.
clear
smaller