Material studied: See herbarium numbers listed in table IV; all material is from East New Guinea (LAEw). Growth rings ar
BLUMEA
22
(1974)
101
—
121
Wood anatomical variation in relation to latitudeanf altitude
van
N.A.
der
Graaff
Rijksherbarium, Leiden,
P. Baas
&
The Netherlands
Contents
Summary
101
Introduction
101
Materials
and methods of this
Limitations
102
study
102
Results
103
Latitudinal
and
altitudinal
Latitudinal
and altitudinal
trends within
103
genera
variation within
108
species
Discussions
109
Infrageneric
trends
109
Infraspecific
variation
114
General conclusions Notes
the
on
115
and
species,
genera,
specimens
studied
116
Acknowledgements
120
References
120
Summary
The wood anatomical
has been
xylem
using
elements
wood
thickenings grouping
scalariform tracheid
to
or
the vessel
helical
anatomical
and
walls.
variation
within
sing
the
to
an
Increasing
for
in
vessel
altitude
2
ray
height with
species
and
and
a
with
or
of secondary
narrower
and
frequency
fibres,
effects,
genera with
and
altitude. An
and 5 with
altitudinal
or
a
wide
none on
vessel
partly or exclusively
altitudinal
wood
Podocarpus
of the wood
analysis
did
range
altitude. The results
variation in
lower
of helical
expression
latitude. In the softwood genus
increasing
and latitude
and
altitude and latitude
to
miniaturization
sometimes
perforation plate in
with altitude
decrease
of latitudinal
wood
latitude
has similar but much weaker
the above mentioned features
phylogenetic
shorter
wide latitudinal
a
in relation
genera
increasing
frequency
The number of bars per
and
hardwood
vessels,
narrower
increase
previous interpretations
significance
eurytherm
generalnot correlated
diameter
reveal correlations between
with reference
with
in
17
from 52 species. With
members,
thickenings.
perforations is
length
samples
vessel
(shorter
reported, together
is
rays)
studied
variation within
are
not
discussed
(Baas, 1973),
stres-
anatomy.
Introduction
In
a
survey
of the wood anatomical
correlation between
species
or
1973).
temperate
perforation,
range
in the
In
regions,
comparing species vessel member
and fibre length decreased with
degree of vessel grouping, and frequency and
*)
present address:
Ethiopia.
genus
Ilex
,
the second author found
a
number of characters and latitude and altitude of provenance of the
involved (Baas,
subtropical per
a
Imperial Ethiopian Government
from the tropics with species
length,
increasing latitude, whilst vessel frequency,
size of bordered
Inst,
from
vessel diameter, number of bars
of
pits
on
Agricultural Research,
the tangential fibre
P.
O. Box
192,
Jimma,
VOL. 22, No.
BLUMEA
102
I, 1974
walls increased. Occurrence and distinct presence of helical thickenings fibre walls
was
for
a
to
material from subtropical
similar but much weaker 'effect'
found with increasing
was
the features helical thickening and fibre pitting. Additional evidence from the literature
Viburnum, and Hydrangea, together with data discussion
general trend in widespread
the vessel and
altitude,
on
montane
except for
Prunus, Symplocos,
(for
rather elaborate
a
an
of
example
a
wood
of relevance for ecological and phylogenetic
genera,
lati-
temperate
tropical
findings for Ilex presented
the
1973) suggested that
Baas,
see
to
to
with those from
tropical lowland
from the
species
tudes. In comparing
regions,
deal restricted great
a
anatomy.
The present to
study
was
undertaken in order
test
(belonging
genera
to
14
mous
genus.
species, This
cal
40
was
In order
specimens
whether
any
studied of
5
genera.
the
on
genus
Wood samples used
level. The results
are
anatomical features The latter
Means and
5
were
square
one
examine
to
gymnosper-
also be traced within eurytherm
can
in
presented
are
AND
or
altitudinal
range.
and ad-
tables, diagrams,
was
ranges
measurements.
Franklin's
to
millimetres. Vessel frequency has are
tangential
not
based
sections and
height
ray
were
counts
macera-
are
1968).
in all
based
determined in
been recorded for ring-porous on
was
diameter. Wood
vessel diameter, number of bars
Vessel frequency and vessel grouping
semi-ring-porous woods the values
5 cm
method (Anonymous,
diameter, fibre length, and
fibre
from other
material studied
tangential, and radial
transverse,
for vessel member length,
Rijksherbarium and
Not all the
branches exceeding
stems or
prepared according
were
METHODS
(1967).
Stern
to
from
studied in
perforation, tangential 25
also done for
wide latitudinal
a
from the wood collection of the
vouched. Almost all material
least
with
was
17
studied in order
was
notes.
wood collections indicated according
per
This
of the trends
species
MATERIALS
tions.
species
52
and
dicotyledonous
considered to be of crucial importance for tracing back the origin of the anatomi-
variation
ditional
to see
were
on more genera
mentionedabove. From
families), the wood of
latitudinal and altitudinal variation within
collect additional data
to
the general validity of the suggestions
representative
on
at
woods; zones
at
least
for
of the
growth ring. LIMITATIONS
Unless stated otherwise, the material studied or
branches with
a
diameter of
at
vessel member length had become this diameter
of
may
have influenced
our
course
occur,
derived from this
For Ilex
cm.
more or
was
STUDY
less stable
and other
gave
peripheral
a
parts
of stems
sufficient guarantee that
(Baas, 1973)- Further changes beyond
sources
of within-a-tree variation
may
also
have influenced results. However,we believe that such variation cannot
the general trends, derived from The botanical
least 5
THIS
OF
identity
many
of the material
species and wood samples. may
unvouched and for the vouched specimens However, the wood anatomical character
be uncertain for those specimens which of
in need of
genera
complex
suggested
a
that
were
taxonomic revision. no
major misidenti-
fications had occurred.
The material studied has mainly been selected wood specimens
in the
collections. This implies that
our
conclusions for the
a
genus
as
can
be
or
criterion
in
data for each individual
expected.
genera,
some
genus
whole, because the number
Only after combining the data on all species and latitudinal trends
on one
Rijksherbarium collection
of
only: the availability of
other institutional
are
too
limited for
species studied
some
was
wood
drawing too
low.
reliability of altitudinal and
N. A.
Ghaaff
van der
& P. Baas: Wood anatomical variation
103
RESULTS
Latitudinal
and
In table I data
altitudinal
given
are
vessel diameter, presence
trends
vessel
on
height for
17
have been
provenance
Latitudinal lowland Zone
I
(L
Zone
II
(L
with
genera
(below
II) Subtropical,
0
(L III)
Zone IV
(L IV) Temperate, 45°
Zone
I
(A
(A
35°
—
0
0 —
45
60°.
—
Zone III
(A III)
—2600
Zone IV
(A IV) 2600—4000 m
zones
regimes
altitudinal
at
m
at
to
23 to
0—500
in table I
m at
to
500 —1200
o° latitude
to
1200
and
I, II, III,
as
IV,
m
as
having
at 23
m at
—2400
coincides
zone
zone
AL
I).
30' latitude
0
23
0
latitude
I, II, III, and IV can be regarded zones
30' latitude (this
0
0 m at
0° latitude
m at o°
1700
Latitudinal
to
L I, and has been referred
1000 —1700
as
alt.):
30'
o° latitude
at
m
zone
II)
II
m
mean
Localities of
range.
zones:
o—1000
with Zone
vessel member length,
fibre diameter, and
follows:
zones as
Warm temperate, 35
I)
fibre length, tangential
1000
0
30'
23
Zone III
solitary vessels, tangential
mean
wide altitudinal and/or latitudinal
into
I) Tropical, o°—23
Tropical altitudinal
percentage of
frequency,
mean
a
grouped
zones
genera
absence of helical thickenings,
or
number of bars per perforation, ray
within
more or
30' latitude
23° 30' latitude.
less similar temperature
for seasonality (Van except
resp.,
Steenis,
1962). Of
specimens
some
the altitudinal
cases
or
data
no exact
latitudinal
the distribution of the species
on
Rhamnus
insufficient
1000
to
In the table
—3000
m
in
given
Kenya.
decide whether
our
they have been included in
zone I. For
can
In order
to compare
character within
a
the data genus
in fractions of 1. In
expressed
on
all
1
and
given the
in
frequency diagrams for
means
plotted
as
i/Vf for
reasons
compared
fig.
with their
studied, to
the
be
per
though
to
a
the
average
I
zone
were
II.
or
the end of this
paper.
Rhamnus and Styrax
can
1,
highest quantitative value for all lower values
having
been
are
given.
Vessel
ray
zones.
height
are
In fig. 3 and
frequency (Vf) has
4
been
of comparison.
1—4
it becomes clear that
tropical relatives,
possess
much lesser
extent,
if
species
more
vessels
one compares
lowland species. The values for number of bars compare
on
perforation, fibre length, and
vessels, shorter vessel members, shorter fibres, and lower present,
notes at
mistakes
the differentaltitudinal and latitudinal
of the relative quantitative values
From table I and
in altitudinal
these values for vessel frequency, vessel diameter,
2
vessel member length, number of bars
distribu-
glabratus
found.
we
genera
in table I, but its
argenteus and S.
the other material more precise data on
possible
has been considered
fig.
to zone I
Styrax
be found under the any
only negatively have influenced the trends
any
assigned
Our data of
will become clear from the results
As
available. In those
in floristic and taxonomic literature. For instance
specimens should be placed
latitude and altitude of provenance
were
had to be chosen somewhat arbitrarily and was based
from East Africa has been
prinoides
tion is from
as
latitude and/or altitude
on
zone
per
from temperate latitudes, if per
rays.
surface
tropical
perforation
area,
narrower
These differences montane
are,
values for different latitudinal and altitudinal
however, zones
in
are
also
with tropical erratic if
fig.
3
we
and
4.
Table
VOL. 22, No.
BLUMEA
104
I.
Variation
distribution. of altitudinal
Species
in
9
wood
Mean values
and latitudinal
anatomical
given
are
in
µm.
zones see
studied
A
s
I
1* §• 5' 5'
text,
N B
B-
(See p.
also
000).
C
