BLUMEA
29
453-479
(1984)
A leaf anatomical contribution to the classification of the
Linaceae complex
P.C. van
Welzen
Rijksherbarium, Leiden,
and P.
Baas
The Netherlands
Summary
The leaf anatomy
riaceae
and
surveyed, mostly
in
of the
Linaceae
Erythroxylaceae)
on
the basis
and
of
complex (Linaceae
allies
putative
Hugoniaceae, Ixonanthaceae,
s.s.,
Humiis
( Ctenolophon, Lepidobotrys, Irvingiaceae)
original observations (72 specimens,
27
genera), partly
from data
the literature.
in stomatal
Diversity
sclereids,
separate family
are
best
to the
or
but leaf
to the
near
thereof
sources
of
in
which
the
the
of the
former.
no
but
The
been
complex. Allantospermum
albeit
more
transferred
clues
strong
its leaf
by
for its true
anatomy
results
some
discussed
in
or
to the
authors.
affinities.
is also in
in
connection
and
near
a
Simarouba-
Lepidobotrys may
than
seems
be
re-
with treatment in
generic
with
of
Cyrillopsis
Ctenolophon
good agreement
deserves
foliar
favour
in
Hugoniaceae
of tracheoidal idioblasts
occurrence
are
treated
closely
Ixonanthaceae, Phyllocosmus
exclusive
evidence
provide
Irvingiaceae (often
complex,
have
they
cells
cristarque
of the Linaceae
Linaceae
Within
account
and
Ixonanthaceae.
the
complex,
the Oxalidaceae.
on
the
anatomy gives
Linaceae
Ochthocosmus
lack
in
with
to
Ixonanthaceae
unrelated,
lated
similarities
petiole vascularisation, sclerenchyma support,
and
cavities,
secretory
of the members
accommodated
show
ceae)
status
midrib
type,
mucilage cells,
status next
in the latter
evidence
from
to
and
other
enquiry.
Introduction
The
Linaceae
sensu
lato
are
currently
being
revised for
Flora
Malesiana
(Kool,
1980; Van Hooren & Nooteboom, 1984) and the present leaf anatomical study
world-wide basis
was
undertaken
as
a
complementary
means
to
on a
provide arguments
in
the debate of taxonomic delimitation above the genus level. Over the years the Linaceae
a
complex
has
been treated very
detailed historical account,
1.
Are the
various
Hugoniaceae,
throxylaceae)
distinct
For
to
suprageneric
taxa
Ixonanthoideae
or
family
views
see
various authors. Without
(Linoideae
or
can
be summarised
Linaceae
s.s.,
going
as
status, and how
into
follows:
Hugonioideae
Ixonanthaceae, and Erythroxyloideae
and various isolated genera
merit
diverging
differently by
the controversial issues
or
or
Ery-
(Ctenolophon, Lepidobotrys) sufficiently
are
Bentham & Hooker
their
(1862),
mutual, phylogenetic affinities?
Hallier
(1923),
Winkler (1931),
454
BLUMEA
Leonard
(1981),
2. Are
not
and
which
3.
Young (in
are
Which
the
are
has
question
treated
at
a
or
(1912,
of the
the
between the
a
moot
as
an
the taxonomic
history
ones,
et
al.,
a
plemented
complex,
and if
(1873),
Oliver
Leonard
(1931),
samples
data from
played
an
groups
the literature
(Rury,
1965; Noote-
Forman
(I.e.)
micromorphological
even
charac-
important role (Solereder,
integral
has
but
species,
since
1899
1913; Winkler, 1931; Metcalfe &
updating.
Erythroxylaceae
and
This
has been
Allantospermum
in the Ixonanthaceae.
an
1903; Boas,
is in need of
and
1965)
Irvingiaceae
(or Geraniales)?
Irvingiaceae (Capuron,
of these alliances,
have
and the
complex
genus
subfamily
1968). However,
(Humiriaceae,
with
the
(Forman,
as
1901; Van Tieghem,
number of
groups
Winkler
(1931),
Linaceae
of the
ally
Irvingioideae
descriptive information
sampled
the Linaceae
to
of the order Rutales
since
point
hand using leaf anatomical data of all
rather low
(1964),
(1981), Cronquist
Compare opinions by
Knuth
1923),
Ixonanthaceae
the
1950; Metcalfe
Scholz
(1959),
Thome
1982).
Simaroubaceae)
leaf anatomical
1908; Jadin,
the
Hutchinson
all related
at
relatives?
living
relationships
to treat
ters, including
Chalk,
1984
Mendonga (1951b).
various authors
by
Throughout
&
closest
become
1967)
proposed
Bedell & Reveal,
Lepidobotrys
Hallier
and Exell &
(or Irvingioideae
boom,
and
their
(1903),
(1950),
2,
(1973), Takhtajan (1980), Dahlgren (1980),
Ctenolophon
Engler
No.
29,
Mendomja (1951a),
Exell &
(1950),
Shaw
Airy
VOL.
-
discussion of the
never
Our present
study
information
and
problems
been given, and much of
on
Simaroubaceae)
1981a &
is restricted
the
b; Boas,
most
to
poorly
could be
com-
1913; Vilhena,
1978).
For
easy
classification
paper, is
reference,
at
given
the
and
Genera studied
number of
Linaceae
the conclusions of this
suprageneric
study
the
following
adopted throughout
Other
treatments
Linoideae in Linaceae s.l.
Hesperolinon
Linum
Radiola
Reinwardtia
Tirpitzia
Anisadenia
Hugonioideae
Hugoniaceae
(5)
this
in the literature
genera)
s.s.
(6)
nomenclature
here.
Family
(total
anticipating
basis of the
Hebepetalum
Hugonia
Indorouchera
Roucheria
Philbornea
in Linaceae s.l.
P.C.
Welzen & P.
van
Baas:
Leaf
of
anatomy
Linaceae
455
s.l.
Ixonanthoideae in Linaceae s.l.
Ixonanthaceae
Cyrillopsis
(5)
Ixonanthes
Ochthocosmus
of Ochthocosmus
Phyllocosmus
subgenus
Allantospermum
in
Lepidobotrys
in Oxalidaceae
Ctenolophon
in Linaceae s.l.
Irvingiaceae
Lepidobotryaceae
(1)
Ctenolophonaceae
(1)
or
Hugoniaceae
Humirioideae in Linaceae s.l.
Humiriaceae
(8)
Humiria
in Linaceae s.l.
Erythroxyloideae
Erythroxylaceae
(4)
Aneulophus
Erythroxylum
Irvingioideae
Irvingiaceae
in Simaroubaceae
or
in Ixonanthaceae
(3)
Desbordesia
Irvingia
Klainedoxa
Simaroubaceae
(c. 20)
Eurycoma
Simarouba
MATERIAL AND
Herbarium material studied is
less stated otherwise
Transverse sections
leaf
margin)
crotome
mainly
(Utrecht: U).
of the
from the
Mature leaves
middle
portion
and
partly
(vols.
sections upon
95
and
: 5)
dehydration
mounted in
in
an
in
peroxide
chloral-lactophenol
The
and
euparal
alcohol series.
glacial
enriched with
studied
specimens
are
were
acetic
together
Aneulophus africana
Anisadenia
saxatilis
Benth.:
Wall.:
Ctenolophon parvifolius
Gabon,
India,
Oliv.:
on a
sledge
paradermal
un-
water.
one
mi-
free hand
safranin/haematoxylin
with unbleached, unstained
in
a
mixture of
volumes of
equal
and unstained leaf clearings
hydrogen peroxide
were
prepared
studied.
listed below:
Nooteboom:
(Capuron)
a
(L)
in
addition Sudan IV-stained cuticu-
Allantospermum
Malaya, FRI6120; Borneo,
—
borneense Forman:
multicaule
prepared
These and
overnight
acid,
Leiden
midrib and
(including
were
stained with
In
at
rehydrated by boiling
petiole
bleached in household bleach.
lar macerations obtained after incubation
30% hydrogen
Rijksherbarium
were
of the lamina
and of distal and basal parts of the
sections of the upper and lower leafsurface
mixture
METHODS
Madagascar, Capuron
Courtet
Hooker
New
&
s.n.
ak
Luang
24560.
(Herb. d'Alleizette).
Thomson
Guinea,
Sibat
SF 23944.
s.n.
Boumann
3355; Philippines, PNH
6385.
A.
456
BLUMEA
Kuhlm.:
Cyrillopsis paraensis
Desbordesia
Durandea
Brazil,
Planchon:
Erythroxylum
Ducke
Pierre:
glaucescens (Engl.)
VOL.
-
No.
29,
1984
2,
Prance et al.
10705,
3783
and Froes
22149.
990.
Zaire, Wagenmans
Hugonia.
see
(Miq.)
cuneatum
Kurz:
S 17894.
Borneo,
E.
-
ecarinatum
Hochr.:
New
Guinea,
BW 9763.
Jack:
Eurycoma longifolia
Fuchs
Borneo,
Smith 2717
Small:
California,
R.Br,
Hugonia afzelii
Lam
car,
Borneo,
&
Schulz
7934
British
(U);
A.C.
Guyana,
Small:
5794.
-
95632.
Sharsmith
4398.
Ivory Coast, Leeuwenberg
2893.
H. costata
Forbes
Miq.: Sumatra,
1010.
Leighton
and
H.
-
California,
H.
-
drymarioides (Cur-
Sharsmith 4162.
Planch.:
ex
Meeuse
SAN 44657
Sleumer
Surinam,
(U).
Hesperolinon adenophyllum (Gray)
ran)
21334.
Benth.:
Hebepetalum humiriifolium (Planch.)
H. jenkinsii
-
Schlechter
racemosa
(Durandea
H. castanea Baill.:
-
2814.
F.v.M.: New
-
H.
cf.
Guinea,
Van
New
deplanchei Stapf):
MadagasMiq.:
costata
Royen
&
Caledonia,
Balansa 2372.
Humiria
I.
St. Hil.: French
balsamifera
Indorouchera
contestiana
griffithiana (Planch.)
Hall, f.:
BE:
Borneo,
BS
Jack:
Burma,
China, How 70738; Borneo,
Klainedoxa
Pierre
gabonensis
Zenker
Olsen
Kostermans
var.
gary,
Filarzky
Ochthocosmus
Cleason:
O.
Desf.:
corymbiferum
barrae
29033.
congolensis (De
O.
Froes
Haviland
Oliv.:
malayana
I. reticulata
-
7932
and S
ex
2951
van
25178
&
Benth.:
De Wild.:
Trabut
Borneo,
SAN
7687.
New
2840.
-
Guinea,
26093.
I.
-
petiolaris
7364;
Pullen
129.
Van
Le
sections
&
42755
Ducke
Testu
1265.
dolomiticum
Borb.:
Hun-
305.
Steyermark
Brazil,
L.
—
Soest
Adderley
Louis 6151.
Zaire,
(paradermal
and
& Hose
den Brink f. 5485.
12066.
Crete,
(ibid.)
Jack:
(isotype); Africa,
&
L.:
Wurdack
roraimae
s.n.
Zenker
Radiola linoides
cult. hort.
& Th.
-
3274
Roth:
cicanoba
Hall, f.:
Klotzsch:
Wild.
Karmann
Cameroun,
lands,
843.
Zenker
94198
I.
-
and
Bakhuizen
7841; Sumatra, Bunnemeijer
Battandier
Venezuela,
magnifolia (Stapf)
Reinwardtia
Brazil,
4034
only).
-
O.
Dunsterville
(paradermal
British
23421;
floribundus
s.n.
(1977).
sections
-
only);
Guyana, Maguire
&
23348.
Phyllocosmus africanus
Zaire,
f.:
-
3328.
L. usitatissimum
-
Hall,
Ducke:
Ducke
Fanshawe
Philbornea
51.
Venezuela, Steyermark
multiflorus
Brazil,
al.
et
Algeria,
(1838).
s.n.
Niel
39257; Java,
oblongifolia Engl,
Lepidobotrys staudtii Engl.: Cameroun,
Linum
SAN
Griffith
528; Philippines,
Van
Borneo,
Borneo,
Irvingia grandifolia Engl.: Cameroun,
Ixonanthes icosandra
Herb. Paris
Guyana,
Hall, f.:
(Pierre)
P.
Dur.)
Germany,
s.n.
5059; Borneo,
Oliv.:
Zaire,
26203.
Gabon,
Dacremont 281.
Courtet
of Ochthocosmus zenkeri
ex
S
Leeuwenberg 4556; Zaire,
Th. & H. Dur.:
Larsen
-
Toroes
Coast,
sessiliflorus
(= type
(Buch.-Ham.
Leiden
Sumatra,
Ivory
s.n.
Karmann
—
(Herb.
s.n.
-
P.
P. dewevrei
Engl.:
d'Alleizette
903);
Hallierf.).
et al. 69.
D.
Don)
R. indica
Hara:
Dum.:
India,
Sri
Hooker & Thomson
Lanka,
Hallier
C
s.n.:
249; Nepal,
the Nether-
Polunin
et
al.
3681.
Roucheria
Columbiana
Buchtien
Simarouba
Tirpitzia
s.n.
-
glauca
sinensis
R.
Hall, f.:
parviflora
DC.:
Columbia,
Ducke:
Florida, Lakela
Hallier:
Tsai
In view of the limited number of
form
leaf anatomical
(tables
1 and
2).
species
-
R. laxiflora
Rio de Janeiro
Long
23423
Winkler:
Bolivia,
(U).
1493.
61751; Tonkin,
AND
descriptions
B.T. 951.
Herb.
29580 and
China, Yunnan,
RESULTS
ed generic
Lehman
Brazil,
Bon
1754/5.
DISCUSSIONS
studied per genus,
but
present
the
we
refrain from detail-
descriptive
data in tabular
Some information not included in the tables will be discussed in
the survey of characters. The leaf anatomical
diversity
in the Linaceae
complex
lends
P.C.
itself well for
Welzen &
van
diagnostic
value of the differences
use
in relation
of
our
but
457
s.l.
material should be studied
more
to
does
the
test
invali-
not
discussion of overall leaf anatomical similarities
a
dif-
or
delimitationand affinities above the
level.
genus
of the leaf anatomical characters with
Survey
Linaceae
Leaf anatomy of
here. This lack of comprehensiveness
taxonomic
to
Baas:
purposes,
reported
data for
date the
ferences
P.
comments
on
diagnostic
and taxonomic
value
Indumentum and papillae
Most
variable below the
(here
cies
(Fig.
studied have
taxa
1981 a
(cf. Rury,
1
a-d)
&
b), Irvingia
inflated,
and
acuminate
tion with
hairs
but
in combination with
unicellular hairs
along
(Metcalfe
hairs).
young
glandular
1950). (5)
in
hairs
Tufted hairs
shoots and floral
also
are
The
parts.
leaves
mature
of
is
very
is
family
to
Epidermal
The
of this
cells
x
8
provide
(cf.
—
Baas
125
cells than
et
varies
x
to
may
be
al.,
be
an
to
see
some
leaves of Cteno-
mature
limited taxonomic value
or
whole
study
of
more
)
above the
Hugonia
perhaps
or
is
spe-
species and specimens
whether the remarkable indumentum of
Although
genus.
appearance)
on
young
absent from
vegetative
and
the
mature
floral
mature
additional argument for the isolated
and
position
and
fairly
in
10
crystalliferous cells)
cells vary
straight
nature
1982:
from
70 pm
trees.
in
(including mucilage
variable
reputedly
Cell size
8
the
(stellate
unspecialised epidermal
the genera appear
groups
for
for Ctenolo-
typical
genus.
ied. Anticlinal walls
the
for
constant
Ctenolophon
status
this and
test
the tufted hairs
of
Further
(Linum, Reinwardtia).
required
Hesperolinon
parts
combina-
reported
and either coincides with sectional delimitation(Hugonia
cies boundaries
leaves,
sev-
with
(numerous
genus level. Below the genus level the variation in Linum, Reinwardtia and
would be
and
glabrous.
indumentum of
interesting
oc-
species). (4) Compara-
Hesperolinon (in
are
(i.e.,
Biseriate hairs
the leaf margin of Reinwardtia indica
Stalked
& Chalk,
to
long top cell),
a
in the other material of this
rare
s.s.),
hairs
prickle
glandular hairs), Hugo-
1950). (3)
with thicker multiseriate stalks
restricted
are
are
The
but
s.s.,
spe-
some
Uniseriate hairs
corymbiferum. (2)
1913; Metcalfe & Chalk,
heads
unicellular
long,
Simaroubaceae
lophon
and in
Irvingiaceae,
number of short basal cells and
a
(Boas,
specimen, exceedingly
glandular
phon
Tirpitzia
Erythroxylum
(1) Simple
occur:
following types
(Linaceae
Hugonia (with
multicellular, glandular
ble
and Desbordesia of the
characterise Linum
cells)
Simaroubaceae
one
in Anisadenia and
Simaroubaceae. Unusual, unicellular
some
in Reinwardtia indica
nia section
in
occur
small, sometimes fairly tall) in Anisadenia and Hesperolinon (Linaceae
Hugonia jenkinsii,
eral
Papillae
of the Linaceae s.s., several
species level)
Simaroubaceae. Trichomes of the
(usually
cur
leaves.
glabrous
to
constant
of
greatly
in size and outline in the
strongly undulating (table
for this feature, which is
anticlinal
epidermal
wall
1).
taxa
As far
surprising
outline in
as
stud-
tested
in view of
many
plant
160).
x
8—146
the abaxial
x
88
pm
epidermis.
in
the adaxial
epidermis,
Herbs and shrubs tend
to
and from
have
larger
458
BLUMEA
1.
Fig.
a-d.
Hair
hair
unicellular
types
in
the Linaceae
of Anisadenia
Hesperolinon drymarioides,
idioblast
of Roucheria
Thin,
taxa,
walls is listed
=
in the
table
in
;
complex,
Palisade
in
stalk
cells
1. They
walls
daughter
develop
are
often associated
cells of the subdivided
into
ly.
Linaceae
ria
species
and Chalk
mucilage
s.s.
and
studied
cells. The
most
by us)
mucilage
cells
(table
1
of
gland.
of Linum
Hugonia
—
e.
division walls
to
be
degree
the
corymbiferum;
d.
costata;
Stalked
b.
Long
gland
of
Subepidermal,inflated cristarque
are
typical
but
for
cells. The
not constant
latter
cells. Undivided
(except
two
in
s.s.
cells also for Roucheria. The
in the
cells
may
also
and the Rouchecells. Metcalfe
Irvingiaceae,
variable for presence
and data from literature cited in the
Hugo-
Periclinal divi-
cells varies great-
mucilage
Hugonia species
are
number of
then the internal
are
epidermal
by mucilaginous epidermal
and Simaroubaceae
a
of anticlinal division
Linaceae
of inflation of the
characterised
common
occurrence
common,
mucilage
epidermal
(1950) reported mucilage
nanthaceae, Erythroxylaceae
of
with
Hugoniaceae
are
hair
of
niaceae and Ixonanthaceae. Their absence is
sion
1984
2,
Prickle hair
a.
epidermis. Only
appear
No.
dotted.
periclinal
or
adaxial
29,
Uniseriate
c.
tracheids
laxiflora.
anticlinal
straight
especially
t
saxatilis
VOL.
-
or
introduction).
Ixo-
absence
Cteno-
P.C.
lophon
of
and
cells
of
groups
character.
Yet
(Linaceae
s.s.
case
Welzen
lack
Lepidobotrys
mucilage
and
van
recognise
can
Hugoniaceae)
and
tary crystals
riera. Some
by
and H.
for
two
us
cells
epidermal
genera
below
the
cells
Boas
species
genus level, presence
taxa
show
infrequent
confined
adaxial
lium is
adnate
The
not
reported
the
to
parallel
ceae
and
most
Tholen & Baas,
balsamifera
of
reder,
here
have
Humiria
to
stomata
an
stomata.
taxa
poles, except
thaceae and
All
a
b).
similar type of
In view of the
In
and Per-
crystalli-
variability
cells
cannot
some
woody
level.
family
or
Only rarely
In the herbaceous genera
region.
abundant all
stomata are
In Linum adaxial
1928)
stomata are
Linum
ap-
tenuifo-
in its leaves which
are
laterocytic
heterogeneous
1931; Vilhena, 1978)
in
at
least
Within the
one
Linaceae
Cyrillopsis
and
to
a
lesser
the
to
Pyykko,
Ter
stomata
subsidiary
in
1962),
1979)
suggestive
to
this genus
cells
some
as
a
in
several
but para-
and Solereder's
of anisocytic
to
stomatal type
anisocytic
stomata
of
separate family.
usually
other
are
(Sole-
special type reported
itself the paracytic
anomocytic
extent
to
sto-
The Hu-
stomata
anisocytic
anomocytic
of Suriana is
the
the
Hartog-Van
stomata.
1950; Smith & Stern,
to treat
stomata, the
Den
to
and
addition
complex
and
anisocytic
an
Lepidobotrys
for stomatal
type. Humiriaceae
genus ( Picramnia;
complex
In
anisocytic
to
cyclocytic
in
7).
according
Simaroubaceae have
additional evidence
paracytic
6 &
Irvingia-
of Phyllocos-
species
embedded in
stomata
anomocytic
to
Erythroxylaceae,
this respect. Two
(terminology
taxonomic marker,
with
in
are
in
(table 1, fig.
has
parallelocytic (2—4 subsidiary
or
Hugoniaceae,
paracytic
cells
of the stomatal
Ctenolophon provide
In the
&
(Luquet,
paracytic
1899; Metcalfe & Chalk,
occur
important
have
balsamifera.
description (1908)
is
a
adaxial
s.s.)
s.s.,
paracytic, anomocytic
(Solereder,
cyclocytic
have
epidermis.
ones.
stomata
constant
1978). Ctenolophon
1899; Winkler,
for
genera
cytic
to
are
neighbouring
miriaceae and Simaroubaceae
record
adaxial
All Linaceae
pore).
paracytic
are
only
Ixonanthaceae
and Humiria
mata
complex.
Hugonia (espe-
cells containing soli-
(Rehfous, 1917; Ozhatay, 1981).
type is predominantly
the
to
cyclocytic pattern
on
(Linaceae
for the genus
have
to
of
stem.
stomatal
cells
mus
constant
species
to
crystalliferous epidermal
in the midrib
the adaxial surface in addition to abaxial
parently
in the Linaceae
epidermal
to
1981
the abaxial
to
stomata
Linum, Radiola and Hesperolinon
over
weakens the
turn
Rigiostachys (= Recchia)
subfamily
at
Linaceae allies
(Figs. 4-12)
mostly
are
known
absence of
or
be used in the discussion of classification
Stomata
recorded
1908; Rury,
value of the
taxonomic
undisputed
cells. This in
confined
are
(1931)
are
absence
or
in this alliance.
occurrence
rare
also
the
mucilage
of the Simaroubaceae:
(Solereder,
The stomatal complex
trend for
Ctenolophon
of
are
459
of presence
variability
limits the
obviously
possess
and
s.l.
(e.g. Hugonia, Erythroxylon, Roucheria)
cells with druses
costata).
Erythroxylum
ferous
to
of Linaceae
anatomy
cells. The
salient
a
Lepidobotrys
Crystalliferous epidermal
afzelii
Leaf
mucilage
related genera
closely
one
the material studied
H.
Baas:
within individual genera
for inclusion of
cially
& P.
do
not
species
touch
at
the
in the Ixonan-
Irvingiaceae.
Hugoniaceae
show
a
remarkable feature of the
subsidiary
cells:
the anticlinal
460
BLUMEA
walls
the
underlying
Slightly similar,
whilst
dolomiticum,
(fig. 9).
For the
cells
guard
but less
Allantospermum
the
Hugoniaceae
No.
29,
sinuous
are
lobes
regular
VOL.
-
(1—6
and
1984
2,
show 3
4
or
subsidiary cell)
per
borneense shows
found
lobe per
one
of the
regular lobing
lobes each
were
subsidiary
(fig.
in
8).
Linum
cell
subsidiary
cells is
highly diag-
nostic.
In
in
vary
less well
developed
spermum the
occur,
lignified
but
The
our
level with
than the
ledges
Hebepetalum,
are
does
not
not
The cuticular
and small shrubs have
two
feature
Hugo-
former genera. In Allanto-
(table 1).
lignified guard cells,
material of Humiria shows
cells
species. Lignified guard
tree
in the
so
a constant
record
epidermis.
species
Roucheria and Philbornea of the
inconspicuously
stomata
the
the herbaceous
The literature
on
but this character is
weakly lignified guard
cells.
hypodermis
A
to
neglected;
in
are
Usually
and inner
outer
Humiriaceae leaf anatomy
often
stomata
for Hugonia. In
also
they
niaceae
the
conspicuousness.
diagnostic
are
section
transverse
ledges
dermal
far
more
common
in the Linaceae
used for classification
The
Hugonia afzelii (Hugoniaceae).
in the midrib region
cells is
development
and
(Ixonanthaceae)
development
lenchymatous
mal
below the entire adaxial surface of the lamina is restricted
complete hypodermis
Ochthocosmus
at
as
a
few translucent
1).
(table
and
complex
Local
parenchymatous
The variation pattern of
allies is such
putative
that it
hypoto
col-
hypoder-
cannot
be
and above the genus level.
mesophyll
Most
studied have
taxa
dorsiventral leaves with adaxial
and abaxial spongy tissue of
layers
Radiola the
of the
mesophyll
leaves.
In
varying
Hesperolinon,
is isobilateral in association with the
and Klainedoxa
Irvingia
(Irvingiaceae)
tissue of 1 —4 cell
palisade
compactness. In
Linum and
condition
amphistomatic
all
mesophyll
cells
are
pali-
sade-like.
In the leaf
margin
the
mesophyll
is
usually
often rather thick-walled isodiametric cells,
cells.
In
palisade
Linum, Radiola, Anisadenia (i.e., all Linaceae except
Hesperolinon,
and Humiria the
zia)
modified and consists of translucent,
sometimes transitional towards
mesophyll
of the leaf
Tirpit-
consists of unmodified chloren-
margin
chyma.
Midrib and
petiole (figs. 2, 17-19)
The range of vascular patterns in the midrib is illustrated
2. The
arc
most
simple type of
of collateral vascular tissue
and
Ctenolophon
has
collateral bundle is
also been recorded
with
strongly
which
ceae,
often
incurved
an
abaxial
p.p.
patterns
with
(Boas,
'pith'
in the
p.p.
margins intergrade
and
arc
1913)
adaxial
and
bundles
are
with
can
be
Humiriaceae
Desbordesia
relatively
simple,
of
rare:
the
s.s.
in
A
figure
simple
p.p., Ixonanthaceae p.p.,
1926; Rury,
Simaroubaceae
'plate'
Lepidobotrys,
diagrammatically
of all Linaceae
Hugoniaceae
(Ballard,
for Picramnia of the
Ixonanthaceae p.p.,
baceae
occurs
Erythroxylaceae
typical
1981
a
&
b).
(Pyykko,
closed vascular
(cf. Colozza,
Ochthocosmus
1904),
Complex
p.p.
Arcs
systems
distinguished (most
Irvingiaceae.
This type
1979).
in
HugoniaSimarouvascular
(Ixonanthaceae),
P.C.
Fig.
2.
van Welzen
of vascular
Types
&
in
systems
P.
Baas:
midrib
and
Xylem hatched; phloem dotted; sclerenchyma
table
1);
a.
continuous
d.
sheath
(marked C).
closed
with
end
—
enclosing
adaxial
Complex
adaxial
closed
bundles
of
a-c.
b.
sclerenchyma support;
e.
—
additional
Erythroxylum
1901; Boas,
superposed
and
ceae
with
parenchyma
with
in
petiole
Simple
s.l.
sclerenchyma;
+ c
bundles
in
1).
table
as
in
S in
with
c.
(marked CCi).
Complex
g.
complex.
systems (marked
(marked
—
461
the Linaceae
open
abaxial
included
(marked CCa).
The
likely
the
on
more
Hebepetalum
—
—
f.
Allantosper-
lack
of
reduction
(cf.
systems
are
provided
matous
ground
vascular
tissue
systems.
systems
also
are
This
a
support
et
like
a
al.,
Linaceae
s.s.
in the
to
the
Irvingia-
borneense the
com-
1982 for
and
'pith'
be taken
derived from
simple
be derived from these
in correlation with the herbace-
midrib of
comparable
parenchyma
as
most
and
Ixonanthaceae
complete cylindrical
the
might
restricted
are
Allantospermum
result of reduction. This is also born
Hugoniaceae
with
main system
In
(cf. Jadin,
systems with adaxial bundles
(fig. 1).
probably
Baas
Within the
might
cylindrical
(Hugoniaceae).
sclerenchyma
and many Simaroubaceae
b)
1924). Complex
small collateral bundles of the
Olacaceae).
cular
less
or
p.p.
subshrub habit is
or
total
1981a &
(Rury,
species
1913; Spiekerkoetter,
is yet of another type
plexity
ous
without
distal
black.
Linaceae
borneense.
mum
one
and
sclerenchyma
closed
Simple
Complex
‘reduced’
Leaf anatomy of
an
closed
Linaceae
argued
the
of
taxa
The
the
open vascular
enclosing parenchy-
with
completely
indication that these
systems.
by
another
reductions in the
simple
fibre sheath,
out
s.s.:
more
simple
closed
open
vas-
complex systems
simple cylindrical (stelar) systems.
It is,
however,
462
BLUMEA
that
likely
increased
complexity
VOL.
nia, and within the species Hebepetalum
character
state
(table
groups
another
to
commonly
is
it
distal part
desia
is
bundles in the
(see
and
open
a
varying
above
(see
under
the
either
(Linaceae
s.s.),
p.p., and the
The
and
or
typically
closely
one
knit
that of the midrib. At the
fibres.
sclerenchyma
At the
halfway
showing
vascular system with adaxial
of
a
complex
the lamina. Desbor-
in the midrib.
are
supporting sclerenchyma.
Apart
are
from
(partial)
a
with collate-
provided
the Linaceae
Only
s.s.
fibres in the othbundle
sclerenchymatous
bundle sheath which is often
of taxonomic interest,
in so-called
(resulting
crystalliferous
p.p.
Hugoniaceae.
or
(cf. Rury,
Fibrous
several Ixonanthaceae
epidermises
1981
a
almost
are
veins),
entirely
linking
the
con-
veins
feature of several
a
bundles
were
found in Anisade-
1), Lepidobotrys,
table
absence of
b), Irvingiaceae, Lepidobotrys,
&
marginal
(cf.
or
or
transcurrent
sheath extensions
both
to
viz., presence
vertically
provided with,
heavily
Vertical bundle
Ctenolophon
Irvingiaceae.
cells
cristarque
crystals
with
lack
only
(Figs, le, 13, 14)
in
and druses
one
the
taxa
studied shows
type represented (table
crystals (table
the
with clustered
(intergrading
1).
usual types
of
solitary,
either in combi-
crystals)
Three genera of the Linaceae
s.s.
1).
Cristarque cells, i.e., crystalliferous
cell wall
within
times
to
with less
closed system
1).
fibres.
crystal complement
rhomboidal
nation
similar
basically
provided
epidermis only,
upper
and Philbornea of the
Crystals
several
Hugo-
the steps from
the midrib pattern
bundle
marginal
Ixonanthaceae, Erythroxylum
nia
occurred
parenchymatous
of the veins
sclerenchyma
the
to
1)
humiriifolium (table
sev-
Judg-
crystals).
continuous
of
Ochthocosmus and
Allantospermum,
secondary and minor veins
vertical bundle sheath extensions
sisting
in
amount
and table
an outer
Two characters
a
parallel specialisations
unrelated assemblages.
support in the midrib also lack sclerenchyma
sclerenchyma
there is
and of
to
simple
a
is
and
identical
dicotyledons
most
veins
sheath,
petiole
exceptional
is
petiole
ral bundles with
er
as
or
and minor veins
Secondary
without
more
virtually
(Irvingiaceae)
As in
1984
2,
1).
The vascular system of the
basis it
have
must
No.
and other related
from the variation within the genera
ing
29,
reduction evolved
or
complex
eral times in the Linaceae
-
(Van Tieghem, 1902, 1903)
cells with
occur
a
thickened and
unilaterally
in various
degrees
of
lignified
conspicuousness
and
in different distribution patterns:
1.
cells
containing solitary
nia,
Hebepetalum,
throxylaceae
very
inconspicuous
of the inner
2.
as
p.p.
This
and restricted
because
dispersed
numerous
in
p.p.,
to
bundle sheath cells in Anisade-
Lepidobotrys,
Ctenolophon
and
type of distribution pattern renders the cristarque
the
sclerenchymatous
1, but also
especially
crystals
Allantospermum
lignified
wall
bundle sheath
throughout
subepidermal
the
portions adjoin
(fig.
ground
layers
the
lignified
Erycells
fibres
14).
tissue of
of the
petiole.
petiole
This
and midrib,
or
distribution pat-
P.C.
Welzen
van
makes the cristarque
tern
Hugoniaceae,
& P.
cells
Phyllocosmus
a
Baas:
Leaf
Linaceae
s.I.
463
feature of the leaf anatomy in
quite striking
p.p.
of
anatomy
(Ixonanthaceae),
most
p.p., and all
Erythroxylaceae
Irvingiaceae (fig. 13).
3.
inflated
cells
cristarque
As
pointed
(Baas,
before
out
than
dicotyledons
druses in the
containing
(Hugoniaceae, fig.
dermis of Roucheria
apparent
cells
1972) cristarque
from the
below the
mesophyll directly
epi-
e).
1
more
are
literature.
descriptive
cells of bundle sheaths have often been overlooked
common
the
Especially
woody
in
cristarque
because of the similar
probably
staining properties of the unilateral wall thickenings and the fibres they adjoin. Their
abundance
nomic
in the
interest.
complex
is
Sclereids
Thinmidrib
more
(Figs.
are
sporadic
difficult
of
to
rare
brachysclereids
and variable
is of great
Irvingiaceae
taxo-
in species of other families of the Linaceae
of these genera
confined
in
occurrence
some
Ixonanthaceae
and
Philbornea)
mus).
none
and
Hugoniaceae
evaluate.
Indorouchera,
In
of
occurrence
15, 16)
thick-walled
to
tissue
ground
Their
they
to
the
of the
ground
tissue of
Hugoniaceae
( Cyrillopsis,
Ochthocos-
Ixonanthes,
constant; sometimes presence
are
and
petiole
(Hebepetalum,
absence
or
even
varies below the species level.
Mesophyll
sclereids
form sclereids
nanthaceae)
occur
and
are
several
Simaroubaceae lack
1901)
was
showed
sporadic,
families. The
and
very
are
of
with
sclereids in
sclerified cells
with the
of
degrees
are
or
of sclereid
(fig.
densely
15).
is
found in
Solereder
Erythroxylaceae (Ballard,
may
as
two
be
doubtful,
constant
a
foliar
be-
feature of
Irvingia species (Jadin,
branching
away
only
from the bundle
sclereids found in the other
thus
unbranched
forming
as
coiled
spiral
(Olatunji
(Solereder,
&
secondary
have
a
three-dimen-
in Roucheria
(Hugo-
genus
are
In
as
these
Spiral-
Xanthophyllum
(Foster,
scattered in the
viz.
spindle-
'eigentiimliche
Ochthocosmus
constant
to
described for
they
1956),
are
and
indepen-
mesophyll together
occurrence
lumping Phyllocosmus
proposed by
(Ixonanthaceae),
referred
1899), Pogonophora
foliar sclereids. Their
as
walls
been
Nengim, 1980).
strong argument against
with the former
Ochthocosmus
(1899) already
dent of the vascular bundles of the veins, and
absent)
other
mesophyll
Our material of Irvingia
branching,
largely
literature similar cells
with 'normal' branched
a
in
records of
latter record
Tieghem (1903).
comparable
1973), Nepenthes
several Orchidaceae
provides
fili-
Eurycoma (Simaroubaceae).
tracheiden'. In the
(Dickison,
not
special type
cells
Van
varying density
tracheoidal idioblasts
shaped
subepidermal
by
also
foliar sclereids
reported
latter show various
sional network
niaceae)
of
are
1923),
The
Irvingiaceae.
inconspicuous
very
these
sheaths, but
A
presence
later contradicted
Slender,
before;
data and literature cited
1904; Hallier,
(Colozza,
(1923) erroneously
and
Irvingiaceae,
(our
In the literature there
and
studied.
taxa
(Hugoniaceae), Ochthocosmus, Phyllocosmus (Ixo-
Simaroubaceae
b)
1981a &
Hallier
cause
value in the
greater diagnostic
sclereids).
sclereids in Humiriaceae
1926; Rury,
of
in Roucheria
(where
in Ochthocosmus
they
Hallier (1923) and Kool
are
(1980).
always
BLUMEA
464
Fig.
4-12.
ensis.
4.
-
Linum
Parallelocytic
sessiliflorus. Paracytic
stomata
Paracytic
to
Paracytic
stomata with
spermum
x
550.
-
laterocytic
borneense.
10.
cells
in
embedded
to
stomata
staudtii.
No.
touching
2,
1984
complex;
at the
anisocytic pattern;
in
with
cells
below
one
lobe
to
x 550.
poles;
x 350.
anisocytic pattern;
Laterocytic
anomocytic
29,
stomatal
Paracytic
lobing of subsidiary
Lepidobotrys
x 350.
VOL.
subsidiary
embedded
stomata
Paracytic
parvifolius. Anisocytic
stomata;
corymbiferum.
stomata with
-
-
x 350.
subsidiary
cell
paracytic stomata; x
stomata; x 550.
-
12.
Cyrillopsis
6.
—
-
8.
350.
balsamifera.
Hugonia
X 550.
beneath
—
para-
Phyllocosmus
7. Humiria
guard cells (arrows);
per
5.
-
X 350.
-
each
11.
9.
castanea.
Allanto-
guard cell;
Ctenolophon
Eurycoma longifolia. Anomocytic
P.C.
Fig.
13-19.
tissue
of
joining
-
13.
petiole;
fibre
mesophyll;
x
van
Indorouchera
x220.
sheath
140.
Irvingia grandifolia.
Welzen & P.
in
-
14.
-
midrib;
16.
griffithiana.
Petiole
with
contestiana.
Midrib
rouchera
griffithiana.
Midrib
Leaf anatomy of
Linaceae
cells
Conspicuous cristarque
Lepidobotrys
staudtii
x550.
Ochthocosmus roraimae.
Phyllocosmus
rouchera
Baas:
-
Foliar
vascular
system
with
thin-walled
central
cells
cavity
in
465
in
peripheral ground
Inconspicuous cristarque
dewevrei.
complex
with
15.
s.l.
sclereids
and
leaf
of
ground tissue;
collapsed
(arrow)
idioblast
140.
clearing; x
mucilage cavities;
central
and traces
in
cell
Tracheoidal
cells
x
x 35.
140.
(f
=
-
-
-
adin
17.
18. Indo19. Indo-
fibres); x
140.
BLUMEA
VOL.
-
29,
No.
£
E o
i-i o
S
3
complement
crystal
2,
I
1984
—
-
-
466
d
d
d
r,
r,
-r,
O
idioblasts
idioblasts
3o
cristarque
-
-
+
-
-
-
_
a
2
tracheoidal
-c CU0 CO Oa
mesophyll
-
-
_
CO
M
sclereids
allies
o 3 3*
i *-• to
—
-
■v.
-
-
33oU
cells
_
CO
CO
cristarque
-
||
5
3 ■*r.
subepidermal
■3
I
S
_
P
—
+
S
_
+
_
—
r
_
+
—
—
Leaf 1.
O
ducts
or
i-i 0
Ho ■*->o
CJ
secretory
5 r3U >,
cavities
-
-
—
-
-
_
co
f-
putative
u
vertically
•a s >
5
3
bundle
and
2).
B
marginal
HI
HC
sclerenchy-
</>a 8
veins
_
ca> |
3 COc
N
transcurrent
-
+
+
-
-
+
±
-
-
<»3
matous
§1
i3S
continuous,
-
+-
midrib
,o
i 'p
table
0c a3 ry.
in
.3 support
oc
*-<
sclerenchyma
"3
a
i
-
+
+
+
±
+
complex and
c
and midrib
'C
a >,
hypodermis
1 ■ac
char cters
data
o3 O
touching
V5
literature
of
U
cells
stomat
CJ
cells
—
D
=a
cells
A
-
-
-
-
-
1
S
S
S
S
(1)
(1) (1)
at
i
subsidiary
jg X> g
I
3M
subsidiary
S
on
c
o nao
lobes
i 3M
6
§
guard
■a
lignified
S
-
-
-
-
-
(i)
(i)
-
CO
O
poles
S
-
-
S
S
S
S
s
s
IS IS
-
-
-
-
-
-
-
-
—
-
—
-
-
-
_s
-
1
-
1
1
-—_+-—_
of
IS
_
see
>
the
S
3
>
vascular
-
fa
II
system
_
Linacea
of
_
petiole
text
a.
type(s)
— in o
i
P
P
P
P
P
P
+
P
P
P
P
P
+
+
+
-
-
P
P
P
P
i
+
—-
mucilage
1 3E
+
+
+
+
+
+
+
_
-
-
-
--
%
-a
_+p - P
+
-
+
+
+
-
-
-
-+P
_
_
-
-
-
-
+
+
+
+
-
-
-
(for
(O
epidermis
Table
>)
Xn
73
adaxially
3 0
K
crystalliferous
common
C0E e 3
£
cells
-
>
\
-
/
-
an tomical
inclusion
o
0
cells
n
A
walls
1i
(
>
>
co
division
n
30
|
outline
c
anticlinal
■a
s-cs-c
3
o
wall
anticlinal
1
studied
Species
s.
s.
s.
s.
Linacea
s-c- s-c- -u- s-u-
u
u
u
u
s-c
s-c
S-f
s-c
saxatils
c-u
c
s-u
u
u
s-c
s-c
sinensi
linoides
HAnisadenia esprolino adenophylum drymariodes Linum corymbiferuLm. dolmitcum usita smiu Radiola Reinwardti cianoba indica Tirpitzia
H. H.
H.
H.
L.
L. L.
L.
R. R.
P.C.
d
d
r,
-r,
_
d
d
d
van
d
Welzen
d
d
r,
r,
r,r,
r,r,
r,r,
r,
I,
r,r,
r,di,
d
-
-
-
-
-
-
-
&
P.
Baas:
d
r,i,
-
d
d
d
d
+
+
+
+
Leaf anatomy of
Linaceae
d
d
d
(d)
r,
r,
r,
r,
-
+
-r,
r
s.l.
467
d/d/-
d
d
r,
-r,d
r,
r,
r,
r,d
d
r,
d
d
r,
-r, -r,
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
+
+
+
-
-
-
-
+
+
+
pages)
next
the
+gt
+gt +gt +gt
+
+gt +gt
+gt
+gt
_+
+gt +gt+gt +gt
-
-
-
-
-
-
-
-
+
±
±
±
+
-
+
-
-
-
+
-
-
_+
- —+-gtP —1—P —ISP+g/t -±— -—+ —P_- P —_+- — —+-—+_
-I± ±n±/? -l±
+
+
CC((C Caa)
_
±gt±gt ±gt+gt gt+gt
+
+
+
+
+c
+c
+
+
c
C
c
c
C
S-C
s-c
c
-1C
+
-
1
-
-
-
-
-
-
-
-
-
-
±
±
±/
±1-
-
-
-
-
-
-
+
+
+
+
C
c
CCi
±
-
-
-
+
+
±
±
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-
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+
±
+
+
+
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+c
s
S
S
s
+c +c
1
-
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s
s
1
1
1C
1
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-
+c— +c—
+
+
-
IS
+c—
+
+
+
+
+
+
+c
c
c
C
c
C
s
S
-1C
-
-
1
CC*
cc*
-
1
1
-
+/+
±/+
1
-/±
-/±
+c— +c—
+c
+c +c +c
s
S
-
S
S
on
(conti ued
IS
1
+
+
+
-
-
±
±
1
f±
/+
1
-
-
-
-
-
-
-
-
-
-
-
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
—
+
+
+
+
+
+
+
+
+
+
-
-
+
+
±
±
±
+
-
-
+
+
-
+/1
+/1
-
-
-
-
-
-
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+/+
-
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-
-
P—
——++
_——
_
P
p
p
-P
—
—
-
+
-
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-l±
-
s-cs-c
u
u
P
p
-
p
p
P
p
p
P
p
-
-
-
-
-
+
+
±
+
±
±
-
-
-
-
+
+
+
+
+
+
+
+
+
-
+
+
+
+
+
+
u
u
s-c
-
u
-
p
humirfolum Hugonia
Durandea
section
section
s-c
+
+
+
+
P
p
-
P
p
P
p
P
p
p
P
p
p
p
pP
p
P
p
-P
-
-
-
-
-
-
-
-
-
-
+/+/-
-
-
-
+
±
±
±
±
-
—
-
-
-
-
-
-
-
-
-
-
-
+/±
+/±
-
-
-
-
-
-
-
+
+
+/+
+/±
±
±
-
-
-
+'
+
-
+
c-uc-u c-c-uu s-c
s-c
s-c
c-u c-u c-u
s
+/—
+/-
s-cs-c s-c
s-c
magnifolai
s
s
±
s-c±
s-c s-c
s-c
(pardemal)
par ensi
thaceae
s-c
P
p
Hugoniace Hebpetalum Hugonia afzeli castanea costa ta Hugonia jenkinsi racemosa Indoruchera contesian grifthian Philbornea Roucheria columbian laxiflora parviflora IxIoxnaothncaen Alantospermu bornense multicaule Cyrilopsi IIxxoonnaatthheess icosandra petiolaris reticulat Ochto smu barae floribundus multiforus
H. H.
H.
H. H.
H. H.
I.
I.
R. R. R.
A. A.
I.
J.
I.
I.
O. O. O.
-
VOL.
d
d
c
E
E o
9
n >. HE3
complement
crystal
r,
s
o3 ■s
cristarque
V, a >> j=aoJ- '-> E
mesophyll
sclereids
3
"s
Vi
VI
3
ducts
•a
or
0
H
Sh
0 +-»
Mu
secretory
1U
cavities
3 c
transcurrent
-3
Is
>>
>
vertically
5
bundle
S3
Ed
g
c
marginal
3|
c
>
veins
.»
g
matous
3
u
u 5/1
o 3S
o
sclerenchycontinuous,
midrib
£.•-
81
oQ. osupport
.s +->
in
E >.
sclerenchyma
x: o cu
S.
petiole
c
E
and
midrib
'C
■3.2
3
2
s
«i vascular
of system
D.
hypodermis
1 o -o o
poles
Si
1
touching
| o 3C
cells
a
[
>>i— .3
subsidiary
'vi£3t«
V>
0
cells
on
C O V*-->
-c
lobes
a 3M •a
c
guard
■a
lignified
8
|
r
r
-
-
d
d
r,
r,
+
+
-
-
±
+
+
),d
d),d
r
-r, -r,
—
(d)
r,(d)
r,
(
-
_
-
+(gt) )
+(gt
(d)
r,(d)
r
-r,
-
-
—
-
-
+
+
—
-+ —P-±+ —-PAi_ + PAi_—- -—+ _ -—±1
_
-
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-
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±
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+
+
+
+
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CCi
C
C
cC
C
c
+/1
-
-
+
C
c C
c
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c
+
+
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c
+
+
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s
s
IS
+
+
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IS
1
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-
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1
1
1
1
1
1
1
1
/-
+/-
±
+
±
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-
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—
—
-
-
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-
-
-
-
-
-
-
±
+
_
-
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+
—-
a
subsidiaryW3
r
-—-
stomat a
v.
cells
=
C4
at
r
+gt+gt +gt+gt +gt+gt
0) 3
VI
u
H
idioblasts
JZ O '■3
tracheoidal
1S
r,
r,
1984
2,
d
d
-
O3
E
Q
|
cristarque
cells
3 o
No.
-r,
VI
c
idioblasts
*-* V,
E C •a'5.5 J= 3 v.
subepidermal
■a
29,
-
BLUMEA
-
468
cells
epidermis
V:
=3.2
■d
M>» i-
O
crystalliferous
cells
3 u
M
o 3E
mucilage
I'
-P -P
PAiPAi PAi
PAi
P
t
P
P
-
-
-
-
_
-P -P
-
-
-
-
-
-
"
_
-
+
+
-
--
-
-
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
_
-
-
-
_
P
-
ou
P
-
>,
X eg c3
adaxially
common
G 0 EE o
Q
f
~
•\
t
P
-
>
i
_
a
type(s)
— Vi
�
>walls
3
c
division
anticlinal
e
.o'v\
|
■3 ■a
1
is
outline
■M 3 oi
wall
■a
anticlinal
|c
+
+
s-c
c-u c-u s-u
s-u c-u c-u
s
s
s-u
s-u
s
s
s-c
s-c
t
roaimae
conti ued)
studied
1
(Table
Species
bals mifera
africana
O(chto smu)Phylocsmu africarnmus congolensi dewevrei se ilforus zenkeri Humirace Humira Erythoxlace AneulophusErythoxylum cuneatum ecarintum
P.
P.
P.
P.
P.
P.
P.
P.
E.
E.
P.C.
(d)
r
r
—
r
r
—
d
- -
r,
r,
r,
r,
van
Welzen
(d)
-
r,
r,
&
P.
Baas:
(d)
-
r,
r,
Leaf anatomy of
Linaceae
d
d
d
d
r,
—
-
(r),(r),
outline:
wall
+gt
+gt+gt +gt+gt +gt
- ---
+
+
±
+
- +_ -
-
+
+
+
+
+
+
+
+
±
+
+
+
_
+
±
_
+
=
=
PAi
+
+
+
+
+
+
+
+
+
+
+
+
+
+
CCa-C
_
CCa CCa CCa
1
1
±1 ±1
±
-
±
±
— —
—
1
- -±1
C
1
1
±/—
±i+c
S
-
+
+
+
+
CCi CCi
1
1
1
1
infrequent. laterocyti ;
poorly
char cter
=
=
L
-
_—
=
C
lobed; CC
=
cell;
per
lobes
3—4
P
P
anomcyti;
=
- - - - P-
cells:
-
P-L
P-L
—
Ai-Aoi-o
-— -—
Ao Ao
Ao
Ao
+
+
+
+
+
—
+
-
±
+
c-u s-u
s-u c-u
-
-
u
u
-
—
-
—
+
c-u
c-u
s
s
s
s
on
char cter
=
Ai
type:
char cter undulating.
=
Irvingacea
gabone si
folia
Desbordesia Irvingia grgranadindifola mal yan Klainedoxa
I.
I.
I.
staudti
parvifoluis
longifolia glglaucaucaa
Lepidobtryace Lepidobtrys Ctenolphace Ctenolphon Simaroubce Eurycoma Simarouba
cells:
forming
and
Ranges
sclernchyma system. solitary
=
C
open;
simple
S=
midrib:
=
+c
of
system
=
u
symbols: straight,
of
only.
region
cells.
midrib
Sclernchyma
2.
fig.
see
suroundi g
special
ar anged
anisocytialy
in
embed ed
curved,
=
=
simple
clusters;
the
e.g.
or
from
adaxialy
as
druses
=
indicated
d
is
bundles,
with
local y
=
1
Hypodermis:
ground below
parenchymatous throughout Varibilty
enclosing present
=
i
bundles, cylinder only;
gt
adaxial complet sheath
=
almost
cytic lobe
bundle
in
a
per
+/—.
species
the
=
with
cell.
ground tis ue.
*
in
=
c
rhomboidal,
r
=
s
Explan tion
vascular
sup ort: open
medulary
pres nt; Stomat l dev loped
+
glauces ns
+
subsidary Vascular type, tissue Crystals: level
absent; anisocytic; Lobes
=
u
u
simple
±
P
a
=
--—
P
com-
i
+
and Ao
—
P
P
complex
=
cy locytic; regular
()
±
-
=
=
dev loped
—
1
P
or
+
one
469
Anticlnal par cytic; iregulary closed; Cristarque crystals.
-—+gt±Ca P —-+ gt - — —-+ ± —
+
+
(+)
(±)
para-
s.l.
or
closed, plete present
hypenatd.
are
states
char cter
transitonal
470
BLUMEA
Secretory
in the form of
structures
extrafloral nectaries
recorded
routinely
is
in
ceae
from the
and
ground
and Philbornea
In
the conspicuous
resin
are
a
several Lina-
epidermis (see above)
cells
mucilage
Hebepetalum they
scattered
Philthe
in
scattered in
even occur
These
they
cavities
ducts of the
'incipient'
but here
they
Chalk
(1950)
but
are
by
no
contain
mucilage.
petiole
and midrib
Met-
(Solereder, 1899;
For Simaroubaceae the
resinous. In Indorouchera p.p., Roucheria parvifolia
as
defined cavities
tissue of the midrib,
mucilage
cavities. Similar
of several Simarouba-
petiole
well-known feature of the
(Hugoniaceae) poorly
ground
the
in
occur
and Metcalfe &
and several Simaroubaceae
Irvingiaceae
down of thin-walled cells.
tory
Noote-
be further discussed.
not
family.
Irvingiaceae
described
p.p.
in the central
or
(1899)
cavities
or
1950).
have been
Lepidobotrys,
will
they
have
p.p.)
In
and/or petiole.
feature of that
canals
calfe & Chalk,
19)
Here
and their pres-
Van Hooren &
(e.g.,
margin,
and also in the Sima-
Hugoniaceae (Hebepetalum, Indorouchera,
Roucheria
Solereder
tissue of the
contents
and
containing oil
to
means a constant
Secretory
the leaf
along
of the lamina.
cells
according
ground
also
doubtfully
mesophyll
ceae
1980).
teeth
complex
by Belin-Depoux (1978)
cells derived from the
Tirpitzia)
tissue of midrib
Secretory
glandular
above under indumentum.
see
mucilage
(Reinwardtia,
s. s.
bornea,
the
hairs
glandular
Apart
1984
the taxonomic literature
boom, 1984; Steyermark & Luteyn,
For
2,
in the Linaceae
widespread
are
roubaceae. Their anatomy has been discussed
ence
No.
29,
(Figs. 17—19)
structures
External secretory
or
VOL.
-
cavities
were
18 &
(figs.
the break-
be considered transitional between
and
Irvingiaceae
probably
observed
possibly originating through
might
were
were
found in the
complete
absence of
peripheral ground
secre-
tissue of
of traumatic origin.
DISCUSSIONS
Taxonomic
In the
implications
following
the leaf anatomical
the genus level. Since
it is
anatomical characters
subfamilies
based
on
families)
or
impossible
alone,
the
will be
Linaceae
and undesirable
surveyed
first. Their
leaf anatomical characters in addition
Linaceae
complex
to
base
a
and their
and above
classification
suprageneric
possible
at
mutual
taxa
on
leaf
(tribes,
relationships
vegetative, pollen and flower
as
mor-
discussed.
s. s.
six
Linaceae genera form
total absence
or
moreover
Tirpitzia
to
subsequently
a
leaf anatomically
paracytic stomata, epidermal mucilage cells,
diola
in the
taxonomic delimitation
to
traditionally recognised
and wood anatomy will be
phology
The
diversity
relatives will be discussed in relation
putative
is
at
least
paucity
share adaxial
exceptional
with
simple
coherent group characterised
of mechanical tissue.
stomata, isobilateral
fairly
well
Hesperolinon,
mesophyll
some
midrib
Linum and Ra-
and absence of
developed sclerenchyma
cular bundle of the midrib. Anisadenia also has
by
vascularisation of the midrib and
crystals.
support of the
sclerenchyma
vas-
and stands
P.C.
within this group
out
features
ler's
&
P.
also
as
of
Leaf
bundle and
marginal
(1923)
Linaceae
of
anatomy
somewhat isolated
a
in Hallier's
implied
(1931) recognition
Baas:
its fibrous
through
be used in support of
can
Linaceae,
Welzen
van
my is
midrib
anatomically
and
chyma
or
(presumably
groups of the Linaceae
and presence of
crystals
(herbs
habit
simple
very
primitive)
more
small
shrubs)
in
the
distinction of 'Anisadenieen' and Wink-
within the Linaceae
Both Anisadenia and
sclerenchyma.
cells. These
of Anisadenia
separate tribe for the genus. Tirpitzia is sometimes said
a
truly intermediate, although
not
471
cristarque
position
be intermediate between Anisadenia and the other Linaceae
to
s.l.
reduced)
The
complex.
more
but its leaf
anato-
Anisadenia shares the
s.s.,
links between the leaf
Tirpitzia provide
and
s.s.,
elaborate
(presumably
absence of scleren-
predominant
leaves in three genera is related
amphistomatic
to
in this group.
Hugoniaceae
The
Hugoniaceae
of lobed
sion
midrib
(only
which
and
in
yet
there is still
Other
stands
a
in
continuous
on
cylinder
to
If the vascular
system
define genera
cells
cristarque
cell walls
epidermal
sections:
or
unlignified guard cells;
The
ground
ill-defined cavities
a
tendency
Ixonanthaceaeand
fined
presumably
form
to
Hugonia
the
by
are
are
leaf
presence
(cf.
tables
hypodermal
vascular
The
be
and
cells
in
(except
systems
simple
complex
interpreted
anti-
break-down of the central
are
2).
rather diverse.
define the
also of
are
two
quite
so
system
as
an
characters
might
be indi-
where
different if
with
as
included
occurs
only
be de-
occurrence
most
outside
Ixonanthaceae
(embedded
in
and
an
aniso-
some
form
unlignified guard cells,
the
subsidiary
cells have
un-
one
cavities. The various midrib and
one
in
could
related groups and
species only)
include
fibre sheaths in
closed system
common
stomata
Phyllocosmus
Allantospermum
not
They
putatively
Positive characters for
and absence of secretory
species)
continuous, cylindrical
reduction of the
can
cells
in one
with
1
development. Negative
subsidiary
lobe each
p.p.
anatomically
of anticlinal division walls, paracytic
cytic pattern of surrounding
tems
lysigenous
presence of characters which
Ixonanthaceae
petiole
undulating
1).
secretory cavities.
absence of characters which
the
lobed
from the
partly
differs from
Allantospermum
Ixonanthaceae
constant
of
resulting
open,
Roucheria
Philbornea has
tissue of the midrib in Indorouchera p.p. and Philborneap.p.
cative of
The
survey of characters and table
(cf.
is
containing druses and its
section Durandea because of its indumentum of uniseriate hairs and
clinal
characters
cells, lignified guard cells,
veins; within the genus Hugonia, section
transcurrent
vertically
help
subepidermal
sheaths);
fibres.
sclerenchyma
characters
and
petiole
bundle
to
group. Variable
mucilage
are
tissue of
restricted
are
of the midrib.
sclereids; Indorouchera has
mesophyll
cells
their shared posses-
by
ground
feature of this
constant
Hugoniaceae
of
defined
strictly
cells in the
cristarque
system
of its
account
very
cristarque
the
the
closed vascular
leaf anatomical
out
and
another
are
predominate
simple
a
cells
Hebepetalum
stomata
paracytic
leaf anatomically
are
subsidiary
considers the
Cyrillopsis
Phyllocosmus
medullary
simple
open sys-
and Ixonanthes
bundles of Ochthocosmus
elaboration on this basic pattern.
as
a
and Ochthocosmus
p.p.
472
BLUMEA
The
its
Cyrillopsis,
see
well leaf
Allantospermum
could both
they
heterogeneity
dence
of
species
two
anatomically
to
(cf.
1984
2,
rather different from each other, but leaf
are
be included in
extent
any great
No.
29,
the Ixonanthaceae without
in the
only recently incorporated
and is
anatomically,
&
(Robson
Airy Shaw, 1962),
leaf anatomical
support.
sentatives of these
close
especially
Ixonanthaceae
Celastraceae and
is
This conclusion
families in
the
based
fits
1980)
assignments
to
would find
Cyrillaceae
comparing slides of repre-
on
reference collection and
Rijksherbarium
evi-
(for supporting
1973, and Kool,
Ixonanthes. Previous
to
increasing
2).
tables 1 and
Forman, 1965; Nooteboom, 1967; Hutchinson,
Irvingiaceae
less
VOL.
-
data
on
from the literature.
The
subdivision of Ixonanthes into
nanthes
Kool
other
(the
finds
(1980)
two
some
species)
I. reticulata
The
and
(1923)
shared
specimen)
again
branched
Kool
by
vide
a
significant
sections Decastemon
and
Phyllocosmus
has
Phyllocosmus
(in
(the
its
epidermal
of
surrounding
in
the
opposition
two
genera
the
major
and maintained
other
cell pattern
Yet
indeed
by
P.
1).
Hallier
by
the
Through
mutually close,
for Ochthocosmus
but the
s.s.
pro-
the
and P.
sessiliflorus
its
absent from
table
(1965).
by
two
zenkeri)
differ somewhat in their leaf anatomy. Section
species)
cells albeit in low
of the
(also
through
both genera apart. Within Phyllocosmus
material represented
our
out
combined
Forman
to
are
been
and exclusive
constant
keep
to
cristarque
cells).
ico sandra) and Ixo-
cells
mucilage
have
Phyllocosmus
frequency
absent from Decastemon. The latter section
are
(I.
(1923)
locally developed hypodermis (cf.
(1980)
idioblasts,
argument
a
and
foliar sclereids
remarkable tracheoidal
of
epidermal cells,
and of
Ochthocosmus
genera
Hallier
support in leaf anatomy: I. icosandra stands
lack of anticlinal divisions in the
one
sections: Brewstera
two
suggested by
as
stomatal
and
moreover
in
complex (paracytic
these differences
are
in
difference between the genera Ochthocosmus and
out
on
account
anisocytic
less
opinion
our
whilst these
indistinct,
stands
of
pattern
than
significant
Phyllocosmus.
Lepidobotryaceae
This
monotypic
families studied,
cytic
stomata
family
has
in addition
to
a
bination of characters is
too
Oxalidaceae
(1950,
by
in
in
or
suggested by
near
the
Leonard
some
without
somewhat
ones.
Other
common
in the
(Hallier,
to
features.
striking
through
its
noteworthy
of the veins and the
Lepidobotrys
several authors
1958) family concept
treatment
(as
quite
'characterless'
out
paracytic
tical bundle sheath extensions of
itself is
leaf anatomy
stands
Lepidobotrys
tendency
characters
cristarque
Ixonanthaceae but leaf
or
to
for lateroare
the
cells. This
oppose claims for closer
1923;Knuth, 1931)
Within the
ver-
com-
anatomy by
affinity
with
support Leonard's
for the genus. Leaf anatomy is neutral with
respect
Oxalidaceae
I.e.).
or
between Linaceae
See Metcalfe & Chalk
(1950)
s.s.
and
for
to
Erythroxylaceae
an account
of Oxali-
daceae leaf anatomy.
Ctenolophonaceae
Ctenolophon
cytic
very
stomata.
common
differs from
the Linaceae
complex through
Other leaf anatomical characters of this
nature
and do
not
allow
conclusions of
its
anomocytic
to
monogeneric family
phylogenetic
affinity.
aniso-
are
of
a
The in-
P.C.
dumentum of the
considered in this
of
alignment
of
tems
in other
single
of
occurrence
these
though
Olacaceae
cells
(Baas
al.,
et
most
as
reflected
in the
classification
recent
sys-
leaf anatomical ranges
larger
1975 and Baas
also separate
tufts)
from all other families
as
complex
Linaceae
order in
(cf. Baas,
might
also known
are
the
same
well
as
473
al., 1982 for diversity
et
Ilex, and the coherent family of the Olacaceae). The sporadic
cristarque
cells
close allies,
its
s.l.
hair
vegetative parts (stellate
in the
natural groups
Linaceae
of
anatomy
be refuted in view of much
cannot
genus
Leaf
Affinities with
study.
doubtlessly
within the
Baas:
Linaceae and
Ctenolophonaceae
angiosperms
P.
and young
calyx
from the
Ctenolophon
Welzen &
van
to
(Jansen
in Celastraceae
occur
families
1982);
be cited in favour of such
even
which
to
Ctenolophon
1973)
and
assigned
once
was
al-
alliance,
an
& Baas,
(Oliver, 187.3; Hallier, 1912, 1923).
Humiriaceae
The
Humiriaceae
leaf
with
a
rather
sibilities
(Vilhena,
the
Ctenolophon;
cytic
and
could be
tendency
some extent
to
of Vantanea
stomata
1899, 1908;
to our
considered in this
data
on
together
towards
in
laterocytic
Humiria
(Metcalfe
the anomocytic
to
to
of
stomata
anisocytic
& Chalk,
1950)
Endopleu-
in Humiriastrum
stomata
link the
family
The
cells
stomata
reminiscent of Lepidobotrys
are
study.
surrounding epidermal
p.p. of the Ixonanthaceae; the
compared
also
taxa
pattern of
anisocytic
an
Phyllocosmus
1978)
addition
in
the range of stomatal types is considerable and
embedded in
stomata
(cf. Solereder,
combinationof other leaf anatomical characters this opens pos-
general
of Humiria recall
1978)
diverse
quite
1950; Vilhena, 1978
advocate affinities with almost all the
to
paracytic
anatomically
Chalk,
species only). Especially
one
ra
are
1904; Metcalfe &
Colozza,
.
of
(Vilhena,
The
para-
the Linaceae
to
complex.
most
In
the
as
of the
family
not
modern systems
order
same
Linaceae.
in conflict with
prehensive study
wards
plex
generic
are
the Humiriaceae
Linaceae. Thome
Leaf anatomy
presumed
of all species
to
treated
are
neutral
is
as
and Melchior
to
a
separate family
(1964)
consider it
either treatment, and
in
a
would
to
doubtlessly
be
rewarding
as
a
the
sub-
certainly
affinities between Humiriaceae and Linaceae. A
characterisation and
closest
(1981)
com-
contribution to-
indicate which members of the Linaceae
com-
the Humiriaceae.
Erythroxylaceae
There
mily
Rury
in
a
s.s.,
diversity
Irvingiaceae,
Mostly
ceae
general
same
(1981a
Linaceae
ing
is
the
&
and
as
the
our
to treat
Erythroxylaceae
Linaceae. Leaf anatomy
results
own
(table
1) fully
which
can
partly
be
Simaroubaceae and
treated
a
as
a
leaf
subfamily
as
separate fa-
a
studied
by
confirm the affinities with
family
there is
ecological adaptations
an
interest-
(Rury, I.e.).
Allantospermum
in the Simaroubaceae, the
anatomically
cells, paracytic
interpreted
as
comprehensively
Ixonanthaceae and Humiriaceae. Within the
constitute
mucilage
b)
of opinion
concensus
order
stomata,
very
mostly
midrib with adaxial bundles in addition
Irvingioideae
coherent group, characterised
to
complex
a
closed
vascular
systems
in
cylinder, vertically
or
Irvingia-
by epidermal
petiole
and
transcurrent
474
BLUMKA
Table 2.
table of
Summary
VOL.
-
important
29,
No.
2,
1984
leaf anatomical characters
and data from the literature cited in the
text.
Legend
based
table
on
1
table 1.
as
CA
=3
8
B
o
M
<D
o
>»
o
a
>>
a
si
type
cells
*CA
J3
X)
3
p
>
o
Linaceae
Mucilage
Stomat l
+
P
s. s.
(A
X)
3
+/-
P
Ixonanthaceae
+/-
P(PAi)
§
Secretory ducts Cristarque
Veins
of
cells
-0)
s
"(+)
_
_
-(+)
+
S-C(-CCa)
+
-(±)
-(±)
+gt(+)
S-C-CCi
+
~(±)
-
-(+gt)
S-CC*
+
S-C
+
+/-
-
-(1)
P
-
£
3
Hugoniaceae
Allantospermum
Sclernchyma midrib transcurent
va
ed
c
caviti
verticaly
u
M
+/±
+/-
-
+/-
P/Ai/Ao/C
Erythroxylaceae
+/-
P
-
S-C(-CCi)
+
Irvingiaceae
+
P
-
(C-)CCa
+
+
+
-
P-L
-
C
+
+
(±)
-
Ai-o
-
s
+
S-C-CCi
+
Humiriaceae
Lepidobotryaceae
Ctenolophonaceae
+/-
Simaroubaceae
cells
cristarque
tomical
ly
diversity
denied. The
rently different
tents
and
ducts in
throughout
absent in
typically
Ao/Ai/C/P
mucilage
veins, conspicuous
-
petiole
and midrib,
petiole
and midrib. This
the Simaroubaceae
s.s.,
but in view
ducts
which
from those of the
characterise
Irvingiaceae
based
tion
of the
1981)
as
my. The
much
on a
distinct from the
of
belong)
or
Capuron (1965),
of
dence
to
the
in
the
of the genus in the
by
Metcalfe
Hutchinson
(1973)
representative sample
Simaroubaceae
assigning
the
family
_7
et
al.
and
Irvingiaceae
as
Ixonanthaceae,
(1968)
and
a
Cronquist (1981).
The
conspicuous
characters
be absolute-
cannot
(1913)
or
are
appa-
fatty
have
by
con-
already
us.
The
or
leaf
anato-
Sapindales (where
Sima-
complex,
to
recogni-
1905; Cronquist,
fully supported by
(1972)
as
discussed below.
have advocated
the
treatment
partly supported by
and
is
ana-
alliance from the Simarouba-
more
opposed
Rojo (1968)
9
+/-
of the rather great leaf
than studied
therefore
view
+
±
and abundant
and Boas
the Rutales
Linaceae alliance is
+gt
-
1886; Van Tieghem,
is
to
-/+(g t)
several Simaroubaceae
(1901)
Nooteboom (1967) and Muller
Allantospermum
(1965)
more
Irvingiaceae (cf. Pierre,
family
problem
roubaceae
-
because of their resinous
also in their ontogeny. Jadin
-
-
combination of
stressed the leaf anatomical distinction of the Irvingia
as
-
in the Simaroubaceae affinities with Irvingiaceae
secretory
possibly
ceae
-
a
position
by
Forman
anatomical
adopted by Airy
Shaw
leaf anatomical evidence
evi-
(1973),
favours the
P.C.
latter
there are
For similar
(1965)
treatment
Shaw
Airy
than
ceae
as
tables
(cf.
The
of
in
leaf
the
cristarque
Ixonanthaceae advocated
the
whilst
to
mainly through
is
cavities of
(?) mucilage
recog-
however, closer
are,
For-
by
implied
does find support
complex
Hugoniaceae
close affinities of
some
the
the
Hugonia-
broad ordinal concept of Geraniales
a
both
including
systems,
with both the Simarouba-
Irvingiaceae
would support
Geraniales
treatment
and arguments from other
combined with
diversity
s.s.,
Linales and
us to
disciplines
alternative
Linaceae
complex
Linaceae
complex
Linaceae
complex
Linaceae
complex
Linaceae
complex
tospermum and
Allan-
(4)
Doubtful
Ctenolophonaceae (1)
Doubtful, Oxalidaceae
(1)
Lepidobotryaceae
family
Cyrillopsis)
(8)
Erythroxylaceae
on
treatment:
Affinity
(5)
(5, including
suggestions
following
favour the
(6)
Ixonanthaceae
Humiriaceae
the
Ixonanthaceae. This
Family (number of genera)
s.s.
profuse
Irvingiaceae,
supported. However,
and the Linaceae
delimitationin the literature have induced
Linaceae
in
be
cells and incipient
complex
anatomical
Hugoniaceae
for the
475
modern systems.
Preferred taxonomic
The
canals and
typical
so
s.l.
should be considered distinct from the Sima-
cannot
to
are
several classical
some
mucilage
Irvingioideae
Irvingiaceae
fairly
and the Linaceae
adopted
the
Linaceae
of
2).
1 and
possibilities
Rutales of
anatomy
and midrib,
(1973)
latter
the
shared, profuse cristarque
ceae
Leaf
In their foliar anatomy the
leaf anatomy.
Irvingiaceae
Baas:
lacks
petiole
as
nition of affinities between
in
P.
why Irvingiaceae
reasons
and
&
differences with Ixonanthaceae.
major
roubaceae, their
man
tissue of
ground
no
Welzen
Allantospermum
treatment.
cells in the
van
or
Linaceae
complex
Linaceae
Irvingiaceae (3)
Simaroubaceae
Wood anatomy
Chalk,
this
distinctly
are
scalariform
to
heterocellular (Kribs
have
(Hugoniaceae
with occasional scalariform
or
complex
large
and
p.p.,
are
exceptional
with
have
not
Rutales,
small,
Tschabold,
vestigially
heterogeneous
half-bordered
necessarily imply
1968)
are
or
both
s.s.
in conflict with
by
or
affinity.
fibres with
The vessel
exclusively simple
plates.
The rays
entirely composed
are
of
or
typierect
Hugonia and Allantospermum
vessel-ray pits.
libriform fibres, but Irvingiaceae share the
1972; Metcalfe &
not
characterised
scalariform
types I—III)
true
is
simple vessel—ray pits.
that of Humiriaceae in essential characters,
cialised, this does
or
Humiriaceae),
cells in the small shrubs and herbs of the Linaceae
resembles
&
1960, and Rojo,
All members of the Linaceae
bordered pits and tend
perforations
cally
1942; Heimsch
Heimsch,
1950; Webber, 1936; Thomas,
treatment.
simple
(cf.
complex
Rutales
(c. 20)
The wood of
but since these
Ctenolophon
are
Simaroubaceae and
large vessel-ray pits
all unspe-
Irvingiaceae
with the
Lina-
476
BLUMEA
VOL.
-
thaceae its wood fits this
The
observations).
better than the
family
wood of
1984
2,
is somewhat apart from the other Ixonan-
complex. Although Allantospermum
ceae
No.
29,
Lepidobotrys
1968 and
Irvingiaceae (Rojo,
known
poorly
too
is
original
the discus-
to enter
sion here.
Pollen
1962 a &
morphology
b)
be
can
1952; Metcalfe
(Erdman,
tospermum have been advocated
al.,
et
different aspects
sing
gy is
for
and Oltmann
1968)
interpreted
both
of
have
to
in
same
the
On
ovules per locule, presence
disk, the
different
which
separable,
awarded
reflected
is
them in all
to
the
Balancing
in
is very
systematic
difficult, partly
biguous arguments
For
support
Linaceae
hand,
s.s.
the
the
an
to
stress
of the
with
complex
suprageneric
the
by
him
of
( Tirpitzia)
with the
might
our
the
more
sults this argument
can
distinct and coherent
than
ing the latter
consequently
finds
number of
extrastaminal
allies
Linaceae
readily
or
family)
multitude of data from
in
account
not
other
analy-
our
provide
unam-
differences between the
groups.
cells
cristarque
(Anisadenia)
and scleren-
be cited in favour of affinities of the
can
and Ixonanthaceae. On
tables
(cf.
number of
a
1 and
separate
be
two
group
also
as
be
as
expanded:
and
Hugoniaceae
treated
a
as
2)
can
the other
s.s.,
Hugo-
be stressed
the
families when he
the
complex,
reasons
the
of
Nooteboom's treatment
support in leaf anatomy is
to
revert
are
as
appear
are
leaf
p.
and differ
recognition
consistency
(1984).
to
The
752).
com-
more
of
and
aberrant
five
our re-
the
most
from the Lina-
(table 2). AcceptHugoniaceae
here
we
and
disagree
which
also
possible family
con-
alternative,
the broadest
as
removed
With
anatomically
and Humiriaceae
Erythroxylaceae
Ixonanthaceae for
also
group' (I.e.
Hugoniaceae
families, requires
Humiriaceae
to
most
families. Each of the
separate
well defined
of the Linaceae
distinct
'Once the
complex):
Ixonanthaceae
reasonably
the latter do from
Van Hooren and
some
are
Cronquist (1981) probably provided
status.
families of the order then appears
with
putative
or
(tribe, subfamily
taken into
or
Erythroxylaceae
recognition of
(=
Linaceae,
that
s.s.
'Celas-
the coherence and distinctness of the Humiriaceae in the order Linales
on
understood
ceae
intra-
an
remaining gaps in leaf anatomical characters between Linaceae
sensible argument for
groups
rank
the
to
has been found similar
(Oltmann, 1971).
and their
either the similarities
support their separate family
from
and
The
status.
petal appendages,
as
cited above, but all
Hugoniaceae,
Ctenolophon
family
treatments.
occurrence
midrib
pollen morpholo-
belonging
as
obturator and
niaceae, Ixonanthaceae and Erythroxylaceae
mented
their
Lepidobotrys
such
Metcalfe
Oltmann all members here
advocate
to
(in
(1972) emphasi-
Simaroubaceae alliance
a
because the leaf anatomy in itself does
instance, the sporadic
chyma
of
leaf anatomical evidence
disciplines (only fragmentarily
sis)
characters
Muller
Irvingiaceae,
and Sarcotheca
Dapania
of the Linaceae
taxa
or
to
and
of
1971; Saad,
Lobreau
by
by
been described
pollen
absence
or
According
pollen
has
and the
combination of various
a
1968; Oltmann,
In these papers the
complex
distinct
that of the Oxalidaceae genera
al.,
Linaceae alliance
a
Cyrillopsis (Ixonanthaceae)
1960),
et
considers that the affinities of Allan-
Irvingiaceae
pollen types.
Linaceae
sufficiently
type' (Thomas,
traceae
and with
and Ixonanthaceae.
families
as
Lepidobotrys
pollen
of the
one
be with Ixonanthaceae
to
(1971)
in favour of either
Irvingiaceae
regarded
if
variously interpreted
P.C.
with
cept
level of the
nomic
and
status
than
does
not
with any
meet
cally
long
as
from this
Ctenolophon
closer leaf anatomical
precise
taxo-
their
as
equal
alliance,
but
can
Metcalfe & Chalk,
or
suggested
in
wood anatomical simila-
1950; Den
Hartog-Van
Ter
al., 1982).
excluded from the Linaceae alliance
be
in the Geraniales close
leaf anatomical,
from the
judged
as
et
treatment
wood anatomical obstacle
the
However,
of interest
in another order for lack of distinctive char-
treatment
complex (cf.
cannot
its leaf anatomy, but
477
recognised.
are
provide
not
1978; Baas
Lepidobotryaceae
Irvingioideae.
hardly
s.l.
candidates. The Celastraceae and Olacaceae
enough
the Linaceae
Tholen & Baas,
Linaceae
Ixonanthoideae, Humirioideae
Hugonioideae,
the
exclusion of
alternatives for
no
Leaf anatomy of
groups is
affinities
favours the
older literature do
rities
Baas:
even
suprageneric
and positive
acters
the
yet
as
possibly
phylogenetic
Leaf anatomy
offer
P.
subfamilies the Linoideae,
as
Erythroxyloideae,
and
Welzen &
van
and
either,
to
be
inclusion in this
common
and
pollen morphological
seems
better match
a
family by
account
on
of
in the Oxalidaceae
to or even
also
probably
macromorphologistudents of
most recent
angiosperm classification.
anatomical evidence
The
Ixonanthaceae, and
Linaceae
complex
should be the
of this
scope
groups
of
could
paper and
angiosperms
as
complicated
very
easily ramify
paper
paper
confirmed
are
Allantospermum
into
the
of 1923
sort
on
by
or
The
of
the
that many of the
acknowledged
of
Rutales,
multidisciplinary study.
Hallier's
in
tive relatives. It should be
but
Irvingiaceae.
for further
subject
treatment
in the
The affinities of the latter with either the
Simaroubaceae and other
the
or
the balance for
tips
in the
not
with both alliances
problem
is
beyond
study embracing
the
many
Linaceae and their puta-
suggestions
in that classic,
the leaf anatomical evidence
presented
here.
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