Ada
801.
Neerl.
Somatic
some
J.C.
December
25(6),
1976,
449-457
p.
chromosome
African
numbers
of
Sapotaceae
Arends
Laboratorium
Plantensystematiek
voor
-geografie, Wageningen
en
SUMMARY
Somatic chromosome numbers of nineteen
ed.
Species
of the
genera
those of Synsepalum
chromosomes.
It is
26 and 2n
=
1.
for the
Autranella, Tieghemella
mon,
2n
(except
=
like
24,
the
,
of African
2n
that
28
=
S.
whereas
26,
=
2n
and
Gambeya
polyploid species
have
concluded
species
Donella
Pachystela,
presents
Sapotaceae have
been determin-
Omphalocarpum
have
subcordatum,
Baillonella
the most
2n
112),
=
toxisperma
2n
28,
=
Tridesmoste-
has
2n
=
24
whereas
primitive number,
polyploid numbers, are secundary.
INTRODUCTION
This paper presents the results of
ling samples, comprising
carried
was
various
(1954) suggested
which
of that
taxa
already
establish the
to
cytologically investigated
ble
at
the
Department
a
study
the
of
Miege
Previously
delimitation of the genera,
offered,
seed-
analysis
seedling morphology
(1977).
have often expressed
of Plant
The
of the chromosome numbers of the
knowledge
to
twenty-four
Sapotaceae.
of the
Bokdam
by
of
analysis
of African
when
a
subject
conflicting opinions.
somatic chromosome numbers of
was
from seeds
produced
karyological
species
family
students of the family
opportunity
were
that
contribute
might
a
with
simultaneously
out
African
Sapotaceae
nineteen
taxa
not
recently seedlings became
and
Taxonomy
collected since the
Geography.
These
on
An
yet
availa-
seedlings
of 1974 in Zaire and
beginning
Cameroon.
2.
MATERIALS
The
taxa
ing
to
investigated
the
two or more
For
in table
presented
Aubreville
samples
with
were
taken
Depending
on
tips
the seedlings
Bokdam’s
specimens
of
their numbers
one to
(1977)
the
arranged
accord-
determination and
paper.
are
herbarium
are
For
five
harvested
seedlings
at
per
various
samples
conserved
shown in
the number of seeds collected
were
column I, and
whereas
some
taxa
have been collected at different times and locations.
also. These
(WAG);
I,
(1964),
identification of the seedlings,
proper
root
are
corresponds
seed
herbarium
tion,
METHODS
of
system
nomenclature
plants
AND
per
sample
in
column
sample
were
and
of the mother
the
IV
on
Wageningen
of
the
their
table.
germina-
analyzed. After analysis
stages of development and subsequent-
450
J. C. ARENDS
The herbarium numbers
ly conserved.
of the conserved
(WAG)
seedlings
are
shown in column 111.
Root
tips
M
20°C,
fixed in
a
a
the
minutes,
root
tips
orcein in
containing 2%
in
in
kept
slightly
sealing
acetic
45%
heated,
with
with
microscope equipped
45%
100
a
hours in
two
karyotypes
and
made
were
The
slide
vapour for
made
9
x
slide.
of
0,5%
then
was
hours,
two
semi-permanent
studied
om
for
tube
test
drop
a
using
oil-immersion apochromatic
x
respectively,
microscope
to a
up in
glass slip.
were
about
at
small
a
in
pretreated
hours
60°C in 1 N HC1
at
acetic acid
blotted,
and
The
a
least
covered with
1,32). Photographs
=
at
five
acetic acid
100%
carefully chopped
were
and
wax.
during
acetic acid, and transferred
45%
firmly pressed
paraffine
objective (N.A.
for
kept
saturated with
petri-dish
a
ethanol and
96%
were
acid,
11.00 p.m. and
to
7°C. After maceration
at
The minute meristematic parts
orcein
10.00
8-hydroxyquinoline
3:1 mixture of
refrigerator
about
at
solution of
aqueous
and stored in
three
collected
were
0,002
12
Carl
a
by
Zeiss
phase-contrast
photographic
cm
plates.
3.
OBSERVATIONS
The method employed
stained
satisfactorily
ultimate stage of
chromosomes,
at
that
one
a
a
metaphase
however,
stage the
slight
of the
stickiness did
mosomes,
correct
a
of
the
the
that,
use
in
of
of stickiness
varying degrees
chromosome numbers
the
Tjio
of
(1948)
is
and
phase-contrast microscopy highly
some
examples
are
=
24, through 2n =26 and 2n
=
=2Bto 2n
24 is found in the
species
number of 2n =26 is found for in total five
respectively
of
Tieghemella,
Synsepalum.
species
(one
species
of
Autranella and
Another twelve
of which
Gambeya:
is
three
most
species
stela. A somatic number of 2n
tum.
while the other
two
species
likely
=
species
=
of
spreading.
but
was
common
generally
fairly
nevertheless for
spreading
aids in
be
out to
In
reserve.
of sticki-
of the chrothe
determining
could be recorded
here
(Jig.
investigation
(column 11,
species, comprising
1
to
5).
vary from
table
have
2n
=
undescribed)
one
and
Tridesmostemon,
finally,
a
1).
The
Baillonella toxisperma. A somatic
of Donella and
112
the
completely,
occurrence
presented
112
general,
The
some
karyotypes
The somatic chromosome numbers found in this
lowest number2n
with
also noticed the
number of chromosomes. Several of the
in
reached
which turned
given
satisfactory staining
from which
are
analysis seriously;
chromosome number
spite
having
distinct when contracted
hamper
not
polyploid species
photographically,
2n
not
species observed, Synsepalum subcordatum,
It is felt
ness.
analysable cells, with,
Chromosomes
contraction often showed the best
were most
occurrence
high polyploid,
analysing
fair number of
a
Since the observed
phenomenon.
low,
yielded
chromosomes.
species
two
28 chromosomes:
of
finally
Omphalocarpum
two
species
of
of
species
three
;
four
Pachy-
is found for Synsepalum subcorda-
Synsepalum investigated
here have 2n
=
26
chromosomes.
Chromosome
morphology
is demonstrated in the
figs.
I
to
5. The overall size
SOMATIC
CHROMOSOME
Provenace
Zaire Zaire
V.
plant WAG
Mother
IV.
NUMBERS OF
SOME
Camero n Zaire Zaire Zaire Zaire Camero n Camero n
4459 4541 4586 4550 4507 4565
4565
4566
Zaire
Zaire
Zaire
Bokdam
Camero n
Zaire Zaire Zaire Zaire Zaire Zairee
Camero n Camero n
82 9-A 82 0-C
8223
7835
Wilde Wilde
Wilde
Wilde Wilde
4402
de de
4553'
4549
E. E.
herbaiurm Bokdam Bokdam Bokdam Bokdam Bokdam Bokdam
451
AFRICAN SAPOTACEAE
J.
J.
J.
4421 4382-A 4564 4392 4312 4567
E.
E.
Bokdam Bokdam Bokdam
F.
F. F,
F,
J.
de
F.
J.
J.
Zaire Zaire
8373-A
s.n.
s.n.
de de
de
Wilde
4457 4420
E,
E, E.
Bokdam Bokdam Bokdam Bokdam Bokdam Bokdam
F. F.
J.
J.
J.
J.
J.
Camero n
de
de
E.
Bokdam Bokdam
F.
J.
J.
J.
17
&
11
Sapotace .
African
specimen
Se dling
III.
111.
4575
Teulings
WAG
4578
1
Bokdam
4631
463
Bokdam
&
&
12
1
4612 4591 4595
16
4630
4630
461
18
4618 4616
4608
15
15
4610 4617
herbaium BBokodakmd4a61m2 Bokdam Bokdam Teulings Bokdam Teulings Bokdam Bokdam Bokdam Teulings Bokdam Bokdam Bokdam
4577,
7
4613
10,
&
8
8
4590 4589 4620
4619
6,
4629
Bokdam Teulings Bokdam Teulings TeulingsBokdam Bokdam Bokdam Bokdam TeulingsBokdam
of
species
2n
IE
II.
26 26 26
112
28
28
28 28
28 28
28
28 28
28
28
28 28
28 28 28 28 28 28 28 26
several
Engler Engler
in
Engl. Engl. Bail on
numbers
(Radik.) (Radik.) (Schum.)
chroms e
Table
De
Engl.
Wild.
ex
Wild.)
(Bak.) De (Pier re
stipulatum stipulatum dulcifcumsubcordaltum bhrevipes bequarti
Somatic
I.
Wild.
Aubrev.
Engl.)
(De
Pel egr.
I.
Pelegr.)
Wild.)
et
(Aubrev.
Engl.
ex
Beauv. Beauv. Beauv. Beauv. Pier e Pier e
(Mildbr.)
(De
(De
P.
P.
P.
P.
P.
P.
procerum procerum procerum procerum lecomteanum lecomteanum spec.
A
Pel egr. Pel egr. Pel egr.
pruniformis ubangiensi spec. beguei lacourtian perpuklchra
spec.
omphalcrpoides
Chev.
A.
Wild.)
(De
—
Pier e PiPier er e
—
congolensi africana ltoxisperma
Sideroxylidea Synsepalum Synsepalum Synsepalum SynsepalumPachystela Pachystela Donela Aubrev. Donela Donela GambeyaAubrev. GambeyaAubrev. GambeyaAubrev. GambeyaOmphalocridea Omphalocrpum Ophalocrpum Omphalocrpum Ophalocrpum Omphalocrpum Ophalocrpum Omphalocrpum Tridesmot n Mimusopidea Autranela TieghemUla Bailonela
et
Taxon
ex
26
26 26
26 24
et
et
et
et
452
J.
1-3.
Figures
orcein
Somatic
chromosomes
staining, phase-contrast,
Figure
1.
Figure
2.
Gambeya beguei
Figure
3,
Gambeya lacourtiana
Synsepalum stipulatum.2n
2n
=
=
28
2n
in
2700
root
tip cells,
=
8-hydroxyquinoline pretreatment,
X.
26
(Seedling Bokdam
(Seedling Teulings
28
C. ARENDS
15
4612
WAG).
WAG).
(Seedling Bokdam
4608
WAG).
SOMATIC
CHROMOSOME NUMBERS OF SOME AFRICAN SAPOTACEAE
Figures
4-5.
orcein
Figure
Figure
cate
4.
5.
chromosomes
toxisperma
2n
root
in
2700
Omphalocarpum procerum
Baillonella
pair
a
Somatic
staining, phase-contrast,
2n
=
=
24
28
(Seedling Teulings
(Seedling
to
mes
vary
are
4.
given karyotype
gradually.
generally
be observed
the
a
One
of the
As
not
can
of
to
a
11
4629
WAG).
WAG).
The
2.5 /im. The relative
species
be noted from the
to
cases
to
appears
Baillonella
complement
distinct. In those
they appeared
karyotype
of
species,
have within its chromosome
(fig. 5, arrows).
tions
Bokdam
arrows
indi-
of longer chromosomes.
chromosomes within
seems to
8-hydroxyquinoline pretreatment,
x.
of the chromosomes varies from about 0,7
and
cells,
tip
453
a
be
length
toxisperma,
pair
of
photographs,
of the
fairly uniform
larger
however,
chromoso-
primary
constric-
where these constrictions could
be median to submedian. Fig. 2 for example shows
Gambeya beguei having (sub)-metacentric chromosomes.
DISCUSSION
It appears from literature that chromosome numbers of the
been
mainly determined
(Mehra
&
Bawa
chromosome
so
1969 and
counts.
far from
Mehra
Further it
can
root
tip
materials.
Sapotaceae
Two
1972), however, published
be concluded that in
only
have
investigators
some
one
meiotic
(Gadella
454
J.
1970)
of thirteen
out
reference is made
publications,
of the materials studied. Since there is much
the
the
materials will
somatic
the
to
24,
=
comparison
Within
one.
is
Pachystela,
According
beya,
represented by
are
world.
=
26
to
have also
number of 2n
three
to
=
28
and
Earlier Tjio
(1948)
too),
was
=
26. The
species
should be
African
placed
of
species
consistently.
of
1970).
The
2n
=
observation
1954,
the
carpum procerum.
which
has 2n
belong
in the
=
to
was
here
a
is
published
However,
in
=
2n
=
=
fig.
26 for
in
=
the
subfamily
had
not
Donella prunifor-
group, Donella
28 chromosomes.
26
chromosomes
28 chromosomes.
=
of Aubre-
opinion
to
Chrysophyllum
accordance
some
other
=
28
with
species
was
of
Omphalo-
28
nevertheless
chromosomes is
presenting
the
karyotype
Tridesmostemon
remaining
Mimusopoideae.
three
to
of
Gadella
clearly
Omphalo-
omphalocarpoides
the
same
subfamily
species investigated
Autranella
found
the results
1962 and
species
(or
oliviforme by Tjio
Mangenot 1957 &
4
yet
chromosome
26 chromosomes for
belong
system of Aubreville (1964) is assigned
26 chromosomes. The
the three
seedling samples, comprising
seven
not
a
this
=
confirm the
to
analyzed
were
the evidence is scanty
have 2n
Gambeya
new
whereas the aforecit-
recorded for C.
seem to
again
2n
the
found 2n
the
to
assigned
Gambeya
species of
have
genus,
for which
Mangenot &
which
chromosome number of 2n
Mangenot &
example
and
(1954)
also have 2n
considered
formerly
present finding of 2n
by
52
next
Donella and Gam-
genera,
species. Although
in separate genera. For the
This
demonstrated
(1954)
S.
the
corroborate the
not
being confined
Gambeya,
two
by
The
stipulatum,
bequartii
species concerned,
(fig. 3)
of
a new
chromosomes,
present investigation
Chrysophyllum
as
two
Miege
remaining
Mihge
Omphalocarpum,
(Miege
next
Gambeya perpulchra,
of Donella and
species
of earlier authors who
carpum
the
American
an
and P.
since Donella and
lacourtiana
and also
number,
whereas the
and
numbers
The somatic
of 26. The
them. These authors
During
data of
brevipes.
the latter
found for the three
species
that
for P.
S.
26
=
multiple
brevipes
of Donella and
cytological findings presented
ville that
four
time.
cainito,
that
appears
multiples of
The
species
Gambeya
Chrysophyllum
P.
by
used
epithets
Chrysophyllum,
ed authors found 2n
ubangiensis
the
(1964),
Gambeya beguei (see /?g. 2)
mis,
of
future. In
distinguished
as
found. This number does
investigated by
that
at
species,
was
the
here,
species,
have 2n
an exact
Chrysophyllum,
the
been
proposed
28
=
the
the other hand is
on
three
by
two
given by Miege(l9s4)
from
Considering
species correctly
been
as
being
two
Aubreville
distinct
Mangenot(1962)
here,
2n
investigation
number of 2n
not
in
four genera have been analyzed.
to
represented
this number
112,
=
specimens
taxonomy
cytological
previous reports.
Sideroxyloideae,
and S. subcordatum. The first
dulcificum
in this
the
species belonging
is
in
subcordatum
Synsepalum
subfamily,
Synsepalum,
genus,
latter 2n
first
of the
published
or
chromosome numbers found
112 for
=
the
several
Aubreville,
first
here,
26 and 28 have all been recorded
number of 2n
it
the
seriously hamper
earlier authors with those presented
respect
2n
the
on
and hence nomenclatural confusion, the lack of reliable refer-
Sapotaceae
ence
herbarium
to
disagreement
ARENDS
C.
congolensis
as
here all
well
as
have both 2n
Tieghemella africana
for these genera have
has 2n
toxisperma
(Aubreville
Aubreville is
Thus it
so
be
can
be
species
finding
concluded that
do
Miege
Further
karyotype morphology
conclusions
provide
geny of
reliable base
a
plant
a
when the
conclusions
towards
on
the
two
having
a
more
even
groups
the
;
of the first
The
species
to
group,
group all have 2n
=
28
of the
cotyledon development
the genera
a
phyloseveral
Sapotaceae
more
seedling-
and Baillonella.
the
chromosomes,
26
here
group,
Donella, Gambeya
advanced
Tieghemella
=
during
analyzed
The first
seedling type.
having
the
concerning
which have been
latter group have lower chromosome numbers of 2n
pectively, irrespective
to
preliminary
of these organs
functioning
according
second
and
endosperm
and
,
species
draw
to
characters which
using
within the
type, includes the genera Synsepalum, Autranella
The
hazardous
more
of
importance
primitive seedling-type comprises
Omphalocarpum
Aubre-
recognized by
as
valid conclusions
the stage of
be
divided into
whether the
data of the earlier authors
cytological
distinguishes
presence
seedling development.
and
the
drawing
subsequent
can
24
by
in the
given
as
to
as
be drawn either. Some cautious and
group. He therefore
seeds and
the
as
be made.
for
types of seedlings based
within
relationship
=
assumed
the evolution of chromosome numbers in the African
can
(1977) points
Bokdam
also recorded 2n
is evident that reliable conclusions in respect
it
cannot
concerning
Sapotaceae, however,
Baillonella
species
Butyrospermum parkii
chromosome numbers
drawing
taken into consideration as well, it would be
are
any conclusions.
(1954)
the
into the subfamilies
taxa
Moreover,
last
to
of similar chromosome numbers.
the
justify
not
arrangement of the various
correct or not.
far. The
so
concluded that
the
supported by
can
26 chromosomes. Chromosome records
=
published
For the latter
it
publication
present
been
not
24 chromosomes and is related
=
1964).
chromosomes,
ville is
455
CHROMOSOME NUMBERS OF SOME AFRICAN SAPOTACEAE
SOMATIC
species
and 2n
=
of the
24
res-
subcordatum (2n
polyploid species Synsepalum
=
112).
Assuming
that there exists
correlation between
a
type and chromosome number, the hypothesis
presents
more
primitive
advanced
having
ling.
a
2n
As
shown earlier,
was
within its
condition in
karyotype.
translocation
to
by
this
These
processes
may have occurred
analyzed by
the
Sapotaceae.
Babcock
two
chromosomes
to
seedling
=
26 and 2n
Baillonella
=
=
28
24
a
toxisperma
advanced type of seed-
relatively long
chromosomes
the product of
may represent
the reduction of chromosome number which
evolution. Similar
(1947) in
Indeed
particularly
has
species
long
leading
during
a
of
primitiveness
be forwarded that 2n
condition and the lower numbers 2n
24 chromosomes possesses
=
can
evolutionary
the well known
have been
processes
example of Crepis
as
referred
Stebbins (1971). Takhtajan(1969) and Hutchinson (1969)
in their
agree
respective
group
works
closely
that
related
the
to
Sapotaceae
should be considered
the Ebenaceae.
Regarding
from the chromosome numbers enumerated
chromosome number of the Ebenaceae
of the
species
having
2n
=
30
by
the latter
Federov
(Diospyros)
chromosomes and
is
advanced
an
family
(1969)
x
some
as
it appears
that the basic
15,
the
having
2n
=
majority
=
60
or
456
2n
J. C. ARENDS
90
=
chromosomes.
Mensbruge
having
the
1966)
far
to
of the Ebenales there appears
numbers which
is
with respect
It should
from its
are
1954),
seedling type, however,
this
different from the
ecologically
for which the
both
found,
less
these
characters.
the
far
supposed
as
As far
Tridesmostemon
already
=
for
Argania
which
regions
Sapotaceae
chromosome number 2n
Bokdam
to
nevertheless
trees
20,
one
the lowest
occur.
investigated species
two
omphalocarpoides
since its
seedling
52,
=
findings
up
show
(2n
=
many
does
26)
characters, including seedling
Tjio
the present
to
by polyploidisation
seems
types. This
1948), Gluena
somes
subcordatum
(table
1)
but also of its
is
not
growth
S. subcordatum is
a
A solution
meters.
an
the
(2n
(2n
28
=
of
was
(personal
primitive
not
meet
are,
type,
as
the
(the
exact
in
only
having
groups
species Chrysophyllum oliviforme
=
48,
here and found
exceptional
problem why
increase in chromosome
Mangenot &
Mangenot
48, Skottsberg 1955). The species
=
species
multiple
of
its
species
of 2n
=
=
112 chromo-
degree
Synsepalum
reaching
tree
this
have 2n
to
regarding
one
dominated rainforest
to
an
occurred
habit. Whereas the other
104 chromosomes
=
to
ivorensis
analyzed
only
indicate,
have
to
pertains
and Planchonella sandvicensis
Synsepalum
2n
that
seedling type. Apart
in which the other
seedling types. According
rainforest
correlation,
the
as
advanced
1962)
of certain
known, primitive.
number
(2n
Also
2n
advanced
chromo-
decreasing
is distributed in arid
species
area
putative primitive
have advanced
communication)
is
Sapotaceae
a
It exhibits many advanced characters. Further the
Pachystela
of
specialisation
be remarked though
species having
a
la
Ebenaceae-Sapotaceae
the forementioned correlation is known:
to
chromosome number found within the
spinosa (Miege
increasing
(De
primitive,
as
evidence it thus
preceding
branch
general tendency
a
correlated with
characters.
morphological
exception
be classified
can
evolutionary
be
from literature
gathered
can
Diospyros
may be concluded that within the
some
be
as
of
foliaceous cotyledons. From the
less
more or
As
seedlings
heights
of
are
ploidy
shrubs,
of about 20
has
2n
26),
the number found
=
112 instead of
so
far for the other species of Synsepalum could be found in the supposition that
,
some
primitive
ancestor
of
Synsepalum
had 2n
=
28 chromosomes. One line of
evolution could have followed the reduction of chromosome
another line
According
to
Mangenot & Mangenot (1962) the flora of the dense
African forest in which the
ancient
ple
2n
origin.
=
polyploid
(1960)
Sapotaceae,
the
diploids.
more
distinguish
numbers,
The
frequent finding
to
a
from which the
of 2n
=
a
the
while
term
was
ancestors are
of the
exam-
by
family
similar level
the
in
as,
levels,
extinct.
Ebenaceae and the
30 in the former
chromosome
for
introduced
plants with apparently
diploid
species
tropical
is of cretaceous, hence
species having
This
link
characterize these
species having higher
occur,
group of
phylogenetic
chromosome numbers of
appropriate
designate
palaeopolyploids.
the forementioned
servation of
appears
as
in order to
chromosome
Considering
species here analyzed
These authors therefore
28 chromosomes
Favarger
number,
developed by polyploidisation.
and the ob-
Sapotaceae,
it
possibly secondary,
such
as
Synsepalum
SOMATIC
CHROMOSOME
subcordatum (2n
that
bility
mesopolyploids
before all
have been
taxa
be
be
species
in its
species
have
be
to
analysis
an
distributional
possi-
designated
be ruled out,
not
Such
thoroughly analyzed.
The
palaeopolyploids.
as
might
but may
unlikely,
of each
samples
designated
polyploid
to
appears
include several
affect
112), could
=
of these
some
457
NUMBERS OF SOME AFRICAN SAPOTACEAE
at
least
as
not
which should
area,
may
again
of the conclusions presented here.
some
ACKNOWLEDGMENT
The author is indebted to his
assistance
in
of Miss Marina
the
growing
is
seedlings
colleagues on the
Wassink in
staff for
preparing
the
critically reading the manuscript.
and of
manuscript
Miss
Maria
The
Teulings
gratefully acknowledged.
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