BLUMEA 23
(1977)
439—447
The secondary
and the
systematic
v.
phloem of
some
position of
Tieghem ex
R.W. den
Ochnaceae
Lophira lanceolata
Keay
Outer
Department of Botany, Agricultural University, Wageningen, The
Netherlands
Summary
The
secondary phloem
of some
representatives
al of
Lophira from the
justified
from
an
ofLophira lanceolata
of the
Ochnoideae
subfamily
anatomical
Tieghem
v.
family, especially
ex
ofthe tribe
Keay (Ochnaceae)
Ochneae of the
(‘Exalbuminosae’),
as
has been
compared with
The
subfamily Ochnoideae.
proposed by
Kanis
(1968),
that
remov-
appears
to
be
standpoint.
Introduction
The Ochnaceae is
in
a
family primarily of shrubs and
tropical Africa, America,
trees
and the East Indies. The
families and five tribes (Kanis,
1968),
viz.
which have
family
can
centres
of distribution
be divided into
two
the subfamily Ochnoideae comprising
sub-
the tribes
Ochneae and Elvasieae Rchb. and the subfamily Sauvagesioideae Lindl. including the tribes
Sauvagesieae, Euthemideae Planch., and Lophireae Rchb. The tribe Ochneae
(Kanis, 1968)
into the subtribes Ochninae and Ouratinae
Sauvagesieae
into
the subtribes
Sauvagesiinae and Luxemburgiinae
(Planch.)
systematic position of the monogeneric African tribe Lophireae, however,
Because of the different
phloem
ofLophira
opinions existing
lanceolata
v.
Tieghem
on
ex
is subdivided
(v. Tieghem) Kanis, and the tribe
this matter,
a
Kanis. The
questionable.
is
comparison of the secondary
Keay with that of
some
representatives
of the
Ochnaceae seemed worthwhile.
MATERIALS
Bark samples
used
from
were
AND
the Versteegh and den Outer collection, Ivory Coast,
West-Africa (1969). All the material studied
from
of small
stems
fixed in FAA.
trees
lanceolata is
were
studied in
macerations. All
sections
Qohansen,
Means of the
1940).
fibre-sclereids
are
based
were
on
designations employed here
and
companion-cell
type
was
vouched. The samples
collected
were
breast height and from basal parts of shrubs, and immediately
at
Only Lophira
Anatomical features
METHODS
at
are
were
a
rather large
transverse,
tree.
radial, and tangential
embedded in Kaiser's
length
least
of sieve-tube
gelatin-glycerin
sections
after
members, parenchyma
twenty-five individual
measurements.
and
treatment
strands,
The
ray
and
type
those of Kribs (1935); the sieve-tube type, sieve-area type,
classified according
to
Zahur (1959).
RESULTS
The results
The axial
are
summarized in table
system of the
1.
conducting secondary
phloem
of the
investigated
Ochnaceae
VOL. 23, No. 2,
BLUMEA
440
species
composed
is
of sieve tubes, companion
1977
cells, parenchyma
cells,
and
sometimes
fibre-sclereids (see plate I and II).
In
Campylospermum flavum,
C.
and
dybowskii,
C.
in
glaberrimum, and
Idertia morsonii,
the main part of the axial system is occupied by parenchyma cells, whereas sieve tubes lie
scattered in between this ground tissue (see plate I, 4). The
transverse
section of the sieve
tubes is almost rectangular and rather small: in tangential direction about
direction about 9 fim. The sieve tubes of Campylospermum
15
flavum, however,
//m,
are
in radial
somewhat
larger.
Ochna
rhizomatosa,
affine, and
Rhabdophyllum
calophyllum
R.
constitute
a
group
inter-
mediate between the first mentioned species and Lophira lanceolata.
In
axial system is
Lophira lanceolata the
constitute 1—3 cell wide (mostly
tial bands that alternate
regularly (see plate II,
Besides
mentioned above;
about 26
The
/um.
7
mentioned in table
the characteristics
those species
sieve tubes and about half of
composed half of
parenchyma cells. They
cell wide) irregular tangen-
one
and 9).
the sieve tubes also differ in size from
I,
tangential direction about
in
radial direction
32 /um, in
section of the sieve tubes is also oval
transverse
round rather than
to
rectangular.
Both
lanceolata
Lophira
and
in Ochna
rhizomatosa,
these fibres
group.
in
occur
viz. about
40
60
jim (see
jum
the
plate II,
(see plate I,
conducting secondary
The crystals occurring in the bark
the first
group,
styloids
Lophira lanceolata;
in
but
intermediate
and
6
group
8). They
are
possess
and 5). Only in Lophira
3
fibre-
somewhat larger
lanceolata
whereas they lie scattered in the species of the intermediate
groups,
While in the intermediate
group the fibres
section of the
in
from
the species
sclereids with a diameter of about
phloem, this
are
isodiametric
species of the intermediate
lanceolata
Lophira
in
about
occupy
is
they
only
to
J
20
percent
of the
cross
Lophira lanceolata.
percent in
somewhat elongated in species of
group,
are
and about isodiametric again
much less
than in the
numerous
other species.
Also
in
stone
cells
occur
in the bark of all the
Lophira lanceolata they
are
The phloem
heterogeneous,
rays
Lophira lanceolata
are
investigated species (see plate I,
often
uniseriate but also
mainly homogeneous phloem
possesses
rays
red-brown
except for
contents,
the axial system sometimes
So the
possess
are
brown
in
a
whereas
seldom uniseriate.
rays
have brown
Lophira lanceolata where only the parenchyma
cells of
contents.
picture produced by the conducting
Lophira lanceolata differs
and 2), only
multiseriate,
which
Both the parenchyma cells of the axial system and those of the phloem
to
1
large quantities.
present in
secondary phloem
near
the cambium of
few points from that of the other investigated species of the
family.
The arrangement of sieve tubes and axial parenchyma
uniseriate
consisting
layers, taking
of parenchyma
scattered within
a
ground
Keay and Rhabdophyllum
irregular tangential
Lophira lanceolata
pound
into
sieve
companion
cells.
that layers
cells is quite often in
affine (Hook, f.)
v.
cells. Only Ochna rhizomatosa (v.
Tieghem
show
a
cells of type
sieve
areas
in
he
Tieghem)
an
phloem. Furthermore
type II with
com-
the tangential walls of the sieve-tube members,
C, fibre-sclereids
consisting of procumbent cells.
species
slight tendency towards
of the elements of the secondary
only species possessing sieve-tube members of
plates, large
tangential
with sieve tubes alternate with those
The sieve tubes of the other investigated
tissue ofparenchyma
layering
is the
account
in
groups,
and homogeneous
phloem
rays
R. W.
Outer:
den
The
Ochnaceae
secondary phloem of some
441
DISCUSSION
The
of the secondary phloem of Lophira differs from all the other investigated
structure
all
species. These species
belong
number of investigated species
Lophira does
is
not
concerned,
Also
the
(Decker,
is
sufficiently known
not
in distinctive metatracheal
parenchyma
and Elvasieae
on
placed Lophira
because of the
compound pitting
between vessel and
It must
in
its
nosae’,
are
and
bands,
Still
homogeneous
abundant
of the fact
that
'
characteristic for the subfamily
highly specialized
with
appear
lacks
Lophira
’
Exalbuminosae (Bailey,
according
to
from all the others,
a
tendency
in broad metatracheal bands in
with the family Ochnaceae it would
rather isolated. This in spite
Also
rays.
and often scanty in the ‘Albumi-
buminosae’ rather than with the ‘Albuminosae’ (Metcalfe and Chalk,
is
distinct tribe,
the tribe Sauvagesieae
structure
more
a
representatives of the
(Sauvagesioideae).
typically paratracheal
occurs
axial
(1966)
absence of unilaterally
like in the other
and its much
Decker
representing
as
pits and the
diffuse in the ‘Exalbuminosae’
it
axial
The tribes Ochneae
rays.
Gilg (1925),
differs in wood
arrangement
however,
affinities of Lophira
‘Albuminosae
family
elements,
diffuse and vasicentric
rays,
following
parenchyma
Lophira
genus
parenchyma
types;
'
subfamily
of the Ochnaceae is
apotracheal,
abaxial aliform
vessel
of
greater diameter
study.
in the
unique
be kept in mind that she studied especially
of the
The parenchyma
several features
of vessel elementswith vestured
Metcalfe and Chalk (1950) the
particularly
subfamily Ochnoideae. "Whether
the anatomy of its secondary phloem
subfamily ‘Exalbuminosae’ (Ochnoideae)
presence
(‘Luxemburgieae’)
as
presents
vasicentric tracheids.
no
in the
‘Exalhuminosae’.
far
the other hand have heterogeneous
and
parenchyma,
as
tracheids,
vasicentric
the
1963, 1969). Although
in the absence of suitable material for
xylem of Lophira
viz.
(Farron,
obvious from the results of this study that
it is
subfamily Sauvagesioideae
secondary
1966),
the tribe Ochneae
small,
fit within the tribe Ochneae of the
quite
fits in the
Lophira
to
is
towards
If there
Lophira.
be with the ‘Exal-
to
1950),
but its
cells
'cristarque'
position
that
are
1933).
'
Kanis
’
nosae
(1968) accepted Engler's (1874)
and ‘Albuminosae’
respectively. The
lack
(Kanis,
1968)
ovuled
ovary
systems
two
distinct subfamilies, named Ochnoideae and
tribe Lophireae
it in the
Dwyer (1941) placed
the seeds
as
shares
it
included by Engler.
not
was
Ochnaceae,
albumen. However,
since
main division of the Ochnaceae into the Exalbumi-
a
more
position
non-distichous leaves
with that subfamily.
in the ‘Exalbuminosae’ because
specifically
in the
Sauvagesioideae
and
a
are not very
secondary phloem,
but
Kanis
may
(1968) but
also
after all also apply
genera
not
because of its anatomy.
to
logical
many-
the
only
This
because of
goes
for
the
the secondary xylem in spite of Decker's
conclusion. Besides this, Metcalfe and Chalk (1950) pointed
of the 'cristarque' cells in the
more
i-celled,
different from each other.
The removal of Lophira from the Ochnoideae appears fully justified,
by
seems
2-carpelled,
Except for the position of the tribe Lophireae,
of Kanis (1968) and Gilg (1925)
the evidence used
Sauvagesioideae
Gilg (1893, 1895, 1925) and
united in the
to
the taxonomic importance
subfamily Ochnoideae
and
Lophira lacks
those cells.
Lophira
it shares
had been
some
vasicentric
tracheids,
parenchyma,
there
may
Lophira.
placed
and
be
also in the
Dipterocarpaceae (Agardh, 1825; Don, 1831)
features with that family, namely
a
metatracheal
axial
cortical bundles,
parenchyma,
chambered
winged exalbuminous seeds. Vestal (1937)
connection
crystalliferous
and Decker
between the Dipterocarpaceae
and
because
vestured vessel
(1966)
pits,
axial
suggest
the Ochnaceae
that
through
BLUMEA
442
VOL. 23, No.
s.a.
2,
1977
simple simple simple simple simple simple simple
7
comp., horizontal, horizontal,
Ochnace
the
horizontal, horizontal, horizontal, horizontal, horizontal,
sieve plates
obi., ±
tube
average member length 7<m
465 220 275 190 190 465 260 320
sieve
type
II
± ± ± ± ± ±
III III III III III III III
of
species
2ui 2ui
bast- type
2ui
s
s
s
s
to
to
s
s
s
6
6
2
some
dbh
with
lanceolat
and
cm
V+O number
20
3
347 388
Lophira
10
64
8
4
10
537
253
and
and
146 798 450 247 785
Far on Far on
Tieghem
of
Thon .) Beauv.) Tieghem
Keay Tieghem Far on Keay
comparison
et
v.
(P.
ex
(Schum.
v.
/.)
Dalz.)
v.
f.)
(Hook,
Tieghem dybowski glaberimu Tieghem) (Hook.
flavum
calophylum
A
1.
et
Table
(Hutch,
v.
studied
lanceolat
Specimen
Lophira
(v.
affine
morsoni rhizomatos
Campylosermu Campylosermu Campylosermu RhabdophylumRhabdophylum
Idertia Ochna
2.
3- 4-
S-
6.
7-
8.
R. W.
den
Outer:
The
secondary phloem ofsome
procumbent;
information
further
ray
maximu number
rays
phloem
of
rows
4
type
phloem-paren- chyma
cells
stone cells
7
5
I-II
Hell Hell
3
3
6
6
II II
6
I
He He He He
4
6
7
8
290 375 260 400 610 580 695
styloid styloid styloid
i:
± ± ± ±
+
+ + + + + + +
groups)
mechanical
companio
3
7
isodiametric isodametric isodametric isodametric isodametric
crystals
tissue
7
Ho
550
/u,m
many most
He
average length strand
of
only
ray ray
II
5—6
of
a
cells cells
II
average number cells per strand
443
upright
crystals upright
few
cells
Ochnaceae
average fibre- sclereid length /rm
cell
type
(in
—
—
A
A
A A
II
II
III III III III III
12
6
6
i.
2.
3-
580
C
type sieve area
Iltg
—
—
960 725 930
A A
A
of
areas
Horizontal diamet r sieve
Species num- ber
side
m
fi
the
on
walls
—
4-
—
—
5- 6.
—
7-
8
VOL.
BLUMEA
444
No.
23,
2,
1977
ACKNOWLEDGEMENTS
Our sincere
Furthermore
and
to
thanks
we
Mrs. T.
are
for Mrs.
I.
Peterkamp-Kuijt
Touw-Koeten who
van
grateful
most
M. T.
to Prof. Dr.
who
prepared
M.
most
of the
who
Willemse,
slides
used
read
critically
for this
the
study.
manuscript,
typed it.
REFERENCES
(1825): Aphorismi Botanici, XVI,
AGARDH, K. A.
BAILEY, I. W.
of vestured
significance
M.
DECKER, J.
in
pits
Wood
(1966):
and
209.
phylogeny
VII.
Arbor.
Arnold
dicotyledons. J.
anatomy
Lundae:
derivative tissues.
and its
The cambium
(1933):
distribution
Structure,
and
diagnostic
14:259—273.
of Luxemburgieae
(Ochnaceae). Phytomorphology
16:39—55.
DON,
G.
D.
ENGLER,
System, I.,
The
(1941):
London:
814.
species
of the
American
Luxemburgieae (Ochnaceae).
Ph.
Fordham
D. Thesis.
New York.
Univ.,
FARRON,
General
(1831):
DWYER, J.
A.
C.
Engl. Bot. Jahrhb.
(1874):
Ochnaceae.
(1963):
Contribution
17:75 —82.
des Ourateae
la taxonomie
a
Engl. (Ochnac6es).
Ber.
Schweiz. Bot. Ges.
73:196—217.
—
(1969):
GILG,
E.
—
Uber
den
nouvelles
(1895):
Ochnaceae.
(1925):
Ochnaceae.
D. A.
JOHANSEN,
du Cameroun
anatomischen
Banks und Tetramerista
Lophira
—
Ochnacees
Deux
(1893):
Miq.
Bau
et
du Gabon.
der Ochnaceae
Andansonia
und
Ber. Deutsch. Bot. Ges.
die
9:115—117.
systematische Stellung
A.
Engler
und K. Prantl:
Die naturlichen
Pflanzenfamilien.
In A.
Engler
und K. Prantl:
Die
Pflanzenfamilien, ed.
In
(1940):
Plant
naturlichen
microtechnique. McGraw-Hill
der
Gattung
11:20
—25.
Book
Company
Leipzig.
2.
Leipzig.
Inc. New York
and London,
523 P-
KANIS,
A.
KRIBS,
D. A.
A revision
(1968):
(1935):
of the Ochnaceae
Salient lines
of structural
of the Indo-Pacific
specialization in
area.
Blumea
the wood
rays
16:1—83.
of
dicotyledons.
Bot.
Gaz.
96:547—557.
METCALFE,
VESTAL,
P.
C. R.,
A.
Hypericaceae
ZAHUR, S.
to
and L. CHALK
(1937):
to
The
(1950): Anatomy
significance
the Guttiferae
Agr. Exp.
St. Mem.
Plate
I.
Ochnaceae.
Tieghem;
2.
Transverse
Idertia morsonii
Plate II. Ochnaceae. Transverse
v.
Tieghem; 7
ca=cambium;
and 9.
cc=
Lophira
companion
et
sections
lanceolata
phloem
v.
TO
of the
THE
Sci.
of 423
in
I. Clarendon
establishing
Press, Oxford.
the
relationship
of the
64:199 —256.
species
of woody
dicotyledons belonging
of the
s=sieve
PLATES
secondary phloem.
Dalz.) Farron;
3
tube;
1
and
4.
Campylospermum dybowskii
and 5. Ochna rhizomatosa
secondary phloem.
Tieghem ex Keay;
cell; cr=crystalliferous
r=phloem-ray parenchyma cell;
anatomy
358, 160 p.
sections
(Hutch,
dicotyledons.
Philipp. J.
of secondary
LEGENDS
v.
of the
comparative
and their allies.
(1959): Comparative study
85 families.
of
8.
6.
(v. Tieghem) Keay.
Rhahdophyllum calophyllum (Hook.f.)
Rhabdophyllum affine (Hook.f.)
v.
Tieghem.
cell; f=fibre-sclereid; p=phloem-parenchyma cell;
sc=stone
cell.
R. W.
den
Outer:
The
secondary phloem of
some
Ochnaceae
445\I
446\II
BLUMEA
VOL.
23,
No.
2,
1977
R. W.
den
Outer:
The
LEGENDS
+
=
=
—
V-fO number
=
dbh
=
type
2ui
=
Outer
den
sieve-tube
=
alternate
sieve
II
=
III
=
are
tubes
small
—3 cell
in
with
or
trees,
wide
diameter
(mostly
tangential
tissue of
ground
Zahur
one
cell
bands
material
of the
Versteegh
of basal
of shrubs
parts
wide) irregular tangentialbands,
of
parenchyma
the
same
width
composed
of
cells
(1959):
with
When the number of sieve
the
closely placed,
very
oblique sieve plates
very
areas
is
plate length
extremely variable,
and the
angle
with
or
10 or
when the
of inclination
were
defining features
as
intermediate
sieve
to
areas.
upon
corresponding herbarium
essentially long (>500 /im),
areas are
relied
1
regularly
scattered
sieve
more
and
cells
according
sieve tubes
height of
constitute
sieve tubes
type, classified
I
sample
collection, Ivory Coast, West-Africa, 1969
diameter at breast
sieve tubes
which
=
TABLE 1
absent
number of the bark
parenchyma
s
TO
447
present
and
bast
Ochnaceae
secondary phloem of some
between
short
are
types
I and
III
with
(100—300 fim)
slightly oblique
to
transverse,
simple
sieve
plates
obi.,
comp., 7
sieve-area
s.a.
type,
=
oblique sieve plate, compound,
classified
II
=
or
Iltg
=
according
a small
a
Zahur
to
number rather
=
small
sieve
areas
(1959):
poorly developed sieve
number rather
sieve
areas on
companion-cell type,
classified
A
the
=
closely placed
areas on
the side
walls, unequally spaced
diffuse
unequally spaced
III
with 7
or
poorly developed sieve
the side walls
according
companion
areas on
the
tangential
side walls
only,
diffuse
to
absent
entirely
Zahur
or
obscure
(1959):
cells
are
much shorter than
cells
are
as
the sieve-tube elements
and
usually
occur
single
C
=
the
companion
strand
of cells
so
that
long
more
as
than
the sieve-tube
one
elements,
companion
cell
but
are
septated to
accompanies
member
phloem-ray
type,
classified
He
=
Ho
=
I
II
=
=
according
to
Kribs
heterogeneousphloem
homogeneousphloem
uniseriate
uniseriate
rays
rays
(1935):
rays;
rays;
procumbent
and
upright cells
only procumbent
and multiseriate
rays
and multiseriate rays
with
long
with short
cells
are
are
present
uniseriate
tails
uniseriate tails
form
a
each sieve-tube
present