Silica grains in woody plants of the neotropics,
especially
B.J.H.
Institute
xylem
of Systematic Botany,
neotropical taxa, especially
examined.
to be
Silica
exist.
grains
tures
in
present
is
(in 80%
value
variable
importance
as a
of the
when
in
a
they
grains,
few
grains
families
They
it
neglected,
is
reported.
very
The
are
were
are restricted to one
often
about 90
most
type
of
characteristic.
great
variation
The
genera).
in
the
secondary
samples
found in the
ray
were
grains proved
paren-
Their dis-
cells).
tissue, although some exceptions
to be
appears
2000
frequently
present
in the
grains
Over
extensively.
and
studied, grains
cases
is
studied
as non-siliceous.
material
silica
Netherlands
and surface texture of silica
were
species (32
far considered
of the
diagnostic
Surinam
300
of siliceous
though,
Size
recognized.
are
so
from
about
especially
very constant,
usually
in
occur
taxa
many
diagnostic
The
is
grains
tissues
chymatous
tribution
The
Distribution patterns, frequency, size, shape,
of
Welle
ter
University of Utrecht,
Summary.
Surinam
in
high.
very
Various
grain
types
size
The
of the
shape
of surface
greatly
struc-
reduces
its
tool.
INTRODUCTION
In 1857
Criiger
dary xylem
of
for the first time described the
some
species
in
what is
balanaceae. The presence of silica
(Klister,
Amos,
1897;
1951,
Murthy,
Some
1952;
investigators
of
Bamber &
have
completely
1960;
Hirata
to
or
the
subject
Burgess,
grains
family
1965;
in the
of the
of many
1932; Prison,
etal., 1972;
dealt with
investigations
1942;
Besson,
1946;
Balan
Menon,
1965;
Scurfield
some
secon-
Chryso-
et
al„ 1974a, 1974b).
evaluate the taxonomical significance
occurrence
of the
technical aspects such
as
of silica grains and the resistance of the
between the presence of silica and difficulties encountered in
saws
made from
grains
by the results of
examined
special steel and sawing under wet conditions, using logs
soaked with water, most
The effect of the silica
The
attempted
1923,
of silica
as
timbers.
By using
judged
Lanyon,
silica, whereas others primarily
timber to marine borers
occurrence
recognized
has since been the
Gonggrijp,
correlation between the
possible
sawing
1903;
1965; Sharma& Rao, 1970;
presence
a
Petrucci,
now
over
a
problems caused by silica inclusions
the resistance
investigations
hundred
diagnostic
on
to
carried out
marine borers is
by
were
only small,
as
solved.
can
be
Southwell & Bultman (1971) who
species.
importance of silica grains,
considered
by
Amos
(1952)
as
a
very
107
B.
J.
H.
TER
WELLE
is still of interest
promising subject,
this
character
tals.
about
Most articles
Asia.
in
as
equal
to
this subject
on
few
a
are
restricted
species
Surinam woods
geographical regions in Africa
(1951) reported the
investigation
20
neotropics,
AND
species
When silica
tropical
of the University
but
Guianas,
samples (c.
samples
family
families,
the Institute of
earlier
of that
backed
are
found in
were
represented
75
on
the
to
the
Chryso-
neotropics; especially
of Utrecht
wood collection of the Institute of
investigated.
were
Besides,
a
great
from Guyana and French Guianawere included in the present study.
grains
genera
included only 44
METHODS
woody species from Surinam represented in the
number of
(1952)
studied because of a good representation in the Utrecht collection.
were
Systematic Botany
Amos
and
of silica grains
occurrence
species belonging
therefore is focussed
MATERIALS
All
to
In his worldwide survey
neotropical species.
with silica inclusions from the
balanaceae. The present
feature like the presence of crys-
comparable
a
Amos
Only Gonggrijp (1923) and
most wood anatomists take the value of
although
that of
c.
a
in
our
Surinam representative
studied
were
wood
as
collection
440 genera and
1300
c.
of
a
family,
well. Most of the families
also
were
species)
the other
not
neo-
found in the
examined. About 2000
were
investigated. All wood
by herbarium material, which is deposited, for the greater part, in
Systematic Botany of
study (ter Welle,
1976) samples
the
University of Utrecht.
for
sectioning
In accordance with
taken from
were
a
an
wood block of
the heartwood.
To
The
the silica
study
sectioning
sectioning
were
rinsed in
water
addition scanning
various
investigation
of 15
electron
types
is restricted
25 /um thick
water
prepared.
were
was
domestic bleaching agent for
microscopy (SEM)
used. After
one or two
the
was
used
to
obtain better infor-
grains.
of siliceous
to
a
to
only cold
and heatedin carbolic acid, and finally mounted in clove oil.
mation about the surface of the
Although
sections
bleached with
the sections
minutes, then
In
distribution, radial
done without any pre-treatment,
was
inclusions
occur
in the
secondary xylem
and distribution of silica
occurrence
this
grains only.
RESULTS
No silica
grains
which silica
observed in the families and genera listed in Table 2.
were
grains
occur
are
grains, the locality and the
frequency,
families
source
and additional data of
are
treated in
cluded. Non-siliceous
tion. The number of
108
described
of the
a
follows:
as
the data
specimens
given species
are
or
on
given in
genus
Species
are
Table
1; size, shape,
reported below. The
alphabetical order. Pertaining data from the literature
genera
of the families
species investigated
is
are
listed
given
at
in
the distribution of the
the end of each family
between brackets.
are
in-
descrip-
SILICA
GRAINS
ANACARDIACEAE
Silica
a
grains
present in all species of Anacardium. Grains
are
maximum size
granular surface,
without a
De Paula&
grain.
de
10
to
one
grain
per
of silica in A.
the material of Loxopterygium
presence
spruceanum. In
contain silica
specimens
grains,
12 -
samples investigated (BBS
6856; BAFOG
198
in Schinus
M). They
with
always globular
a
others do
were
were
sagotii
or
cells
about the
anything
absent in
Hall 57- LBB10734 ; Lindeman
v.
smooth
not. Grains
smooth
a
but there are also
did not report
Hamburgo Alves(1973)
with
globular
cell,
some
wood
of fourteen
eight
5841,6057,6080,6194,
present in the samples listed in Table 1. Grains nearly
size 5
surface,
(1), Spondias (1), Tapirira (2),
lOftm.
to
No silica
grains
were
observed
Thyrsodium (2).
and
BOMBACACEAE
The genus Bombax
Grains
large
B.
5
with
globular
23 pm,
as
siliceous
as
to
into
a
size
If the
genera,
15 fim. The
grains
grains
occur more
were
globular
frequently
all
to
of
without silica
Amos
(1952),
in the
grains;
who
cells
ray
split
and
siliceous
less smooth
as
who studied
Robyns (1963),
grains,
a more or
grains.
but sometimes
comprise
genera
with
15
to
Species
According
contain silica
species. All species of Quararibea
Grains
up
cell.
without silica
species
classification
followed
is
with and
normally
and B. surinamense.
non-siliceous.
is
several
species
but not in every
cell,
per
in the axial parenchyma.
No silica
both
granular surface,
globosum,
species, Bombax
Bombax
comprises
grain
one
B.
flaviflorum,
s.l.
surface,
non-
well
as
size up
in the axial parenchyma than in the ray cells.
observed in Catostemma (2), Ceiba (1), Matisia (1), and Scleronema
(1).
BONNETIACEAE
The
family concept
followed here is
samples
of
globular
and sometimes
two
globular grains
in the
3
to
procumbent
Haploclathra
measure
3
occurrence
studied.
found:
(2)
genera,
are
to
viz.
they
15
ray
De
grains
(1) globular,
irregular,
the
Haploclathra.
other
shapes,
but
sporadically
Paula
are
with
grains
no
Archytea
30
also in the
is
(1974)
noticed silica
in
in
only
one
smooth surface and
look like
a
cluster
Baretta-Kuipers (1976) who studied the
anatomical characters between the
same
grains
compact
occur
Archytea. They
but considered its
types of grains
structure
of small silica
are
size of the
specimens of Kielmeyera
Two
in all
ray cells. The grains in
Haploclathra
specimen.
a
occur
Normally they
X
upright
taxonomic value. Three
the
smooth,
and their surface is less smooth than in
present
a
In
Silica grains
the surface
theoval/oblongonesupto
cells,
in this genus of
Silica
and
Archytea
assume
mostly globular
13
that of Maguire (1972).
(size
up
particles glued
to
8
were
were
/im);
together.
wood samples reports differences in other
sample with and
the
samples without silica.
109
B.
J.
H.
TER
No silica
WELLE
grains
and Neblinaria
were
observed in Bonnetia
Mahurea (2), Marila (2),
(3), Caraipa (3),
(1).
BURSERACEAE
In
this
silica
family
Dacryodes
measure
up
to
m,
except
oval
the
a
presented.
The non-siliceous
glabrescens,
usually
as
in P.
silica
more or
or
less
insigne, they
ones
25
P.
with
a
P.
are;
12ft m.
P.
In
globular
cells, although
with
grains
in
often restricted
a
in her wood anatomical
Canarium
to
granular surface,
the
marginal
size
study
11
to
they
cells
P.
are
a
grains
(1941)
did
and
m.
P.
sometimes
Grains
sometimes,
cluster of small
occur
(Plate 1/4). Grains
22 ftm. Webber
ft
are re-
silica is Trattinickia.
not
of the Burseraceae. No silica grains
(1), Hemicrepidospermum (1),
grains
crassifolium,
present,
look like
occur
up to 17
18
but
up to
ft m,
containing
ray
ones
found in all tissues.
too, the distribution pattern is very variable, but in all species
globular
globular grains
without silica
are
some cases
the
aracouchini,
size
granular surface,
The fourth genus
the
when
they
fibres
cells,
variable. Grains in
Paraprotium the grains
species
grains,
sometimes
m.
In
shapes,
alstonii,
The
pullei.
do not exceed 5
ft
particles glued together.
parenchyma
granular surface;
a
X
with and
species
parenchyma only;
globular,
to
up
species
and
axial
cells,
ray
Grains of various
which both
hostmannii,
restricted to rays
the
distribution pattern is very
oblong with
or
oblong
Protium is
P.
Their
the vessels.
genus in
in
occur
vessels.
globular,
are
in all tissues
grains
of the
and/or tyloses
Here,
in the
ray
more or
less
record silica
were
observed
Tetragastris (4).
CARYOCARACEAE
Silica
grains
globular,
are
size
present
up to
rays. No silica
in
all
13 pm. The
grains
were
species
observed in
investigated
in the axial
grains
of
Anthodiscus.
Grains
nearly
parenchyma resemble those in
the
Caryocar (6).
CHRYSOBALANACEAE
The
occurrence
by Criiger (1857)
of silica
in
some
grains
species
pattern of silica grains in this
tigated
contain
vestigation:
silica
Acioa,
(Plate
in the
family
1/1-2).
Chrysobalanus
,
Parinari.
110
secondary xylem
of this
family.
was
The
made
by
ter
following
Couepia,
was
reported
A worldwide survey
Welle
genera
(1976).
were
Exellodendron
,
for the first time
on
the distribution
All
species
included in
Hirtella,
investhe in-
Licania
and
,
SILICA
GRAINS
CONNARACEAE
Silica
grains
23 p
to
Metcalfe & Chalk
& Veenendaal
m.
The
(1950),
C.
Connarus,
Amos
(unpublished
griffonianus,
(1952)
results)
grains
nor
all
by
Dickison
of
samples
with
globular,
in this
grains
family
is
a
granular
(1972). According
to
Mennega
from Cameroon show the
Agelaea
of this genus,
neotropical species
observed in Cnestidium
were
sur-
reported by
not
Pseudoconnarus and Rourea. An african
as
unlike the
No silica
the ray cells.
of silica
occurrence
silica distribution pattern
same
in
in Pseudoconnarus and Rourea. Grains
occur
size up
face,
and
(1)
of
species
contains silica
Connarus (6).
ERYTHROXYLACEAE
Out of nine species
no
silica
cells and
in
citrifolium; ;
this
smooth surface may
grains
are
size is
15
grains
side
shape
with
side.
by
50
to
globular,
oval
x
20
a
granular
grain
ray
per cell.
reported (Record
Brazier &
In
&
and
in E.
m), except
the literature the
tropical species
a
or
The presence of silica
sample
from New Guinea
may
granular, except
with
grains
less
a more or
highly variable,
one or
two
but there is
of silica
occurrence
in which
shapes
but the
citrifolium where the maximum
absent,
1943; Metcalfe & Chalk,
Franklin, 1961).
as
may be present
grains
Hess,
oblong.
the size is
Consequently
/u
one
All
often determined by the size and
are
Each of the procumbent ray cells contains
cells,
or
may fill up the entire cell. The surface is
they
occur
are
Their size and
species grains
always large (up
right marginal
one
observed. The
sample.
same
of the ray
shape
in E.
grains
in the
occur
of Erythroxylon investigated, E. squamatum is the only
were
never more
in this
grains
in the
grains;
family
up-
than
is not
1950;Normand, 1950; Amos, 1952;
grains is perhaps
(E. ecarinatum)
restricted
studied
by
the
to
me,
neo-
is also
devoid of silica.
EUPHORBIACEAE
In
is
three
genera, Actinostemon,
a constant
the
same
feature. Actinostemon and
characteristic
the entire ray cell
The
In
a
oblong grains
Maprounea they
of the
occur
silica
lumen,
are
are
and
Senefeldera
Senefeldera
distribution pattern.
have
more
a
granular
frequent
small (about
cluster of small silica
grains
Maprounea,
The
surface and
than the
two
are
the
allied genera which share
silica grains, which often fill
they
measure
up
to
3/rm) and of various shapes.
particles glued together. Species
up
from Surinam the grains
to
23 /am, size of the
are
very small
(3
60
x
25
globular grains.
Sometimes they
with and
species
jn Micrandra. Grains globular, with agranular surface, oval
globular grains
of silica
occurrence
oblong
In
ones
two
up
to
species,
or
30
x
M.
glabra
suggest
without silica
oblong,
size
13 pm. In M. elata
and M. spru-
111
B.
J.
H.
ceana
TER
WELLE
silica is
no
present.
These two
species,
formerly
included in the genus
Cunuria,
transferred to Micrandra by Schultes (1952). Webster (1975) reinstated Cunuria.
were
The silica distribution
No
silica
Amanoa
grains
seems
feature in favour for
a
observed
were
in
the
keeping
two
(1), Alchornea (1),
Acalypha
genera apart.
Alchorneopsis (1),
(1),.Aparisthmium'(I). Chaetocarpus (\), Conceveibai(2). Croton (12), Drypetes (2),
Fluggea (1), Glycydendron (1),
Manihot
Hevea
(3),
Hura (1),
Hyeronima (1), Jatropha (1),
Pausandra
(1), Margaritaria (1), Micrandropsis { 1), Omphalea (1),
(4), Piranhea (1), Plukenetia -(1), Sagotia
Phyllanthus
(1),
and
Mabea (3),
Pera
(1),
(2),
Sapium (2).
FLACOURTIACEAE
The wood anatomy of
who
silica
reported
the Flacourtiaceae
grains in the
ray
cells
was
Lindackeria laurina. All other genera from the neotropics
vation is in agreement with my
studied by Miller (1975),
extensively
of Mayna amazonica (size
lack silica
investigation of species
own
10
to
grains.
16
pm)
and
This obser-
from the Guianas.
GUTTIFERAE
Only
2
a
one
10 ,um,
to
of
species
Clusia,
2
mostly
(1974).
In the
granular
not
surface
3
(size
surface
mostly
Oedematopus
pirn)
are
grains.
Grains
smooth. Sometimes the
particles glued together.
of
sample
contains silica
palmicida
5 pm;
to
cluster of small silica
C.
Grains
quadratus
were not
grains
look like
reported by
de Paula
small silica
investigated
present. An unidentified
globular, small,
sample of the
grains
same
with
a
genus did
contain silica.
In fourteen
lack
species
rubella).
species
silica
Grains
up to 25 pm,
uniseriate
(T.
belong
were
investigated silica grains
T.
obovata,
than
According
checked
to
species
by
one
a
per ray
de
Paula
T.
pittieri,
(1974)
the
rileyi,
and
T.
size 3 to 15 pm, sometimes
the
Probably
T.
T.plumeri,
granular surface,
cell.
whereas six other
occur,
grains
occurrence
do
not occur
of silica
in the
grains
is
a
of Tovomita. The herbarium vouchers of the non-siliceous
Mrs. A. R. A.
Gdrts-van
Rijn.
There is
no
doubt that
they
to this genus.
No silica
onia
choisyana,
more
constant feature of all
species
Tovomita
normally globular with
never
rays.
of
(1),
grains
Rheedia
were
observed
in
(3), Symphonia (1),
Calophyllum (1),
and
Havetia
(1), Moronobea (1),
Plat-
Thysanostemon (1).
HIPPOCRATEACEAE
The
wood
genera
112
anatomy
investigated
of
the
this
family
monotypic
was
described
by Mennega (1972).
genus Prionostemma
was
the
only
one
Of the twelve
in which silica
SILICA
grains
found.
were
but other shapes
23
to
This is
pm.
No
silica
in
observed
were
(1),
and Tontelea
(9),
the ray cells and mostly show
to
a
globular shape
well. Their surface is granular and they
as
of the few lianas known sofar
(1), Hemiangium
Salacia
(3),
restricted
are
present
one
grains
Elachyptera
mera
They
are
Anthodon
Hippocratea
to
silica
possess
(2), Hylenea
measure
Cuervea
(4),
Peritassa
(1),
up
grains.
Cheiloclinium
(1),
GRAINS
(1),
Pristi-
(3),
(2).
HUMIRIACEAE
Only
of
species
one
surface,
size of the
there is
without
a
more
In
grain.
No silica
and
than
13
to
in both ray
grains
by Cuatrecasas( 1961).
117 and
(BBS
sometimes up
grain
one
of silica
occurrence
grains
Vantanea
described
sometimes
cell,
per
to
cells
grains
in this
family
Grains with agranuof other
Krukoff 6653)
18
in the axial
amazonica and S.
species (S.
two
1950; Amos,
Chalk,
3
globular grains
observed. The
&
species
mostly globular,
never
contains silica
guianensis,
cells. No difference in silica grain distribution was noted in the
formas and varieties of this
lar
S.
Sacoglottis,
and axial parenchyma
oval
20
up to
ones
parenchyma
cydonioides)
there
15,um;
also cells
are
silica
no
reported
was not
shapes;
x
grains
were
before(Metcalfe
1952).
observed in:
were
Humiria (1), Humiriastrum (1), Schistostemon (1),
(2).
LAURACEAE
Silica grains
Licaria three
out
L.
silica
ones
L.
globular
X
lar surface and
per cell.
ray
in
cells
more
less
and O.
normally
each
contrast,
a
No
The
martiana, L.
with
only
occur
1/5).
it
the
not
aurea,
grains
are
they
are
noticed in M.
splendens,
contains
twenty-six species
were
observed
3
measure
in
grains
one
of
to
and L.
Here the
grain.
The
are
(10),
grains
vernicosa).
with
than
grains
to
are
species
measure
oblong
a
granu-
one
Amos
with-
(1952)
globular
of
are
5
grain
procumbent
although
same
according
be with-
25 pm,
to
in the fibres. The fibres
Ocotea,
Aniba
2
subdivided
13 ,um. Two
to
L. cayennense,
to
about the
pattern,
synandra.
found
never more
in
cluster
a
In the genus
canella,
ones
globular (3
grouped together
This distribution
contain silica
5
were
L.
granular surface, globular
surface and
compartment
(ca.
species
in part of the ray cells. There is
much.
was
smooth
grains
a
small
multiflorum, L.polyphylla,
In these cell families their size is
varies
Nectandra (4), and
aritu,
are
twelve
grains;
look like
Cryptocarya they
grains
L. armeniacum, L.
In Mezilaurus itauba
samples
further
silica
L.
oblong,
Mezilaurus,
or
glaucinia
amara, L.
25
some
(Plate
to
to
they
sample
one
unique
a
In
genera. In
studied have silica
guianensis,
up to 45
neotropical
particles glued together.
species
grains (L.
debilis.
Grains
in four
occur
of small silica
with
Ocotea,
O.
septate and
to
15 fim. In
without silica.
Beilschmiedia
(1), Endlicheria (3),
Systemonodaphne (1).
113
B.
J.
H.
TER
WELLE
LEGUMINOSAE
Silica
grains
family.
with
in the ray
occur
In Dialium
guianense they
size up
granular surface,
a
contain
ferent from that in Dialium.
cells. Grains
axial
silica
with
a
other
No silica
there
size
28/x
m
and parenchyma
in the rays
as
well
in the
marginal cells.
three
grains
per cell.
S.
without silica
and
grains,
S.
are
Four
species
22
jumin
S.
of
contain
globular
guianense.
melinonii and S. micropetalum.
few other genera from the
a
to
up
in the
as
Normally
albiflorum and S. guianense,
albiflorum
cells
Dicorynia is quite dif-
cells
sometimes in the axial parenchyma. Grains
15
in S.
up to
pm
neotropics
contain
silica,
were
observed in Alexa (2), Aldina (1), Anadenanthera (1), Andira (6),
Caesalpinia (1), Calliandra (1), Campsiandra (1), Cassia (2), Cedrelinga (1),
Crudia
Clathropis (1), Copaifera (1),
Derris
or
them,
in
grains
only
occur
of this
all parenchyma
20,um;
both in ray
size up to
two
are
to
of the
occur
genera
parenchyma. Grains globular
Tachigalia.
grains
Bowdichia (1),
grains
Two of
cells and
ray
species,
and
the
the axial
pattern
grains
three
parenchyma cells in
to
sometimes up
granular surface,
Koeppen (1967)
to
Apuleia
a
the rays
granular surface,
According
15
to
investigated.
were
in the
grains
restricted
Here the
but sometimes
one,
The two
e.g.
In
parenchyma.
there is
with
globular
Sclerolobium
are
each. The distribution
grain
one
cells and/or axial
(2), Cynometra (2),
Dahlstedtia
(4), Dimorphandra (2), Diplotropis (2), Dipteryx (4),
Etaballia (1),
(2), Eperua (3), Erythrina (1),
(1), Hymenolobium (2),
Lecointea (1),
Martiodendron
Marmaroxylon (1),
Peltogyne (2), Piptadenia (2),
(1), Pterocarpus
(4),
Samanea
Lonchocarpus (9),
(1),
Piscidia
(1),
Gliricidia (1),
(2),
Mora
Enterolobium
(1), Hymenaea
Machaerium(3), Macrolobium (4),
Pithecellobium(9),
Sesbania (1),
(2),
Heterostemon
Ormosia
(2),
(1), Dalbergia (5),
Elizabetha
Palovea
(5),
Stryphnodendron
Tephrosia (1), Vatairea (1), Vataireopsis (1), Vouacapoua
(1),
Parkia
(2),
Platymiscium (1), Poecilanthe
(1),
and
(3),
Swartzia
(5),
Zygia (2).
LECYTHIDACEAE
The
silica
grains
in
the
frequency. They mostly
in the
one
to
occurs
contain all
sporadically,
grains
All
in
are
in the
rays and
The
in
sample,
The cells
investigated
of
Grains
grains
a
are
grains
or
a
great variation in shape,
sporadically
in
a
one
in Allantoma
are
Cariniana contain silica
grains only,
the
oblong
grain
size
and
parenchyma
and
oblong.
Sometimes
only
or
whole genus, but often samples appear
contain
mostly globular,
size,
in the axial
globular, oval,
seen; in addition
up to 20 /u m, size of the
globular
are
species
normally
grains
found. The
parenchyma.
contain
a
shapes.
globular grains
114
in
cells with two
always
species
the
occur
of the vessels.
tyloses
shape
show
Lecythidaceae
grain,
a
but
globular
grains
and
is 3
to
up to 35
13 pm\
measure
in the rays
sometimes oval
ones
sometimes, although
certain number of cells without
X
or
oblong,
20 p m;
small
up to
m.
and sometimes
in
size
samples
grains
of the
which
often show
a
SILICA
smooth surface unlike the bigger
axial
parenchyma
Corythophora
p
m
in
size.
is
frequency
Grains
ones
even
The
silica
20 ,«m
to
to
up
40
Couratari
they
are
sometimes oval
(partly
30
to
up
in Couratari
larger,
are
more
ray
and
the ray
to
abundant and sometimes they
or
the
20 fi m, and in C. stellata
X
restricted
are
oblong,
oblong
measure
{Krukoff8893 ) up to
granular,
more
oblong
80
are
are
to
up
30
m.
x
and less
x
in the
same
up
to
Their
cells.
sporadic.
size of the
always large,
ones
grains
cells of the
globular with granular surface,
are
in
grains
The
granular.
smaller than those in the
are
grains
sometimes
variable,
mostly globular,
up
grains
the
the surface of which is
ones
pyriformis
In
sample.
13
of C.
GRAINS
globular
15/um
Generally
or
the
than those
frequent
in Cariniana.
Thirty species of Eschweilera
E.
tiflora,
poiteaui,
and
simiorum, four samples
E.
were
In four
analysed.
were
roroda
silica
grains
were
two of which
analysed,
never
in the rays, whereas the other two lacked silica
gated
of
oval
globular,
oblong,
or
oblong grains mostly
up
to
In
50
60
15 ,um in
X
25
size of the
10 /im
x
two
20
/tm)
a
large
globular
30
x
of
ones
20 pm,
to
4
one
to contain
grains.
amount
chartacea,
All other
in
grains
E. congesE.
species,
sporadic
species
the rays.
silica
investiGrains
20 p m, sometimes 25 to 30 p m,
sporadically
up
40
to
15/xmoreven
x
of E. obversa.
globular
but
observed,
were
to
samples
Holopyxidium jaranum
X
show
however,
E.
found. In
appeared
grains
Eschweilera,
species;
grains (up
H.
18
to
and
latifolium {IANw 3879)
oblong grains (up
the
on
to
contrary lacks
silica.
Six
of
species
mostly oval,
globular
are
square
ones
usually
up
is
are
to
18
restricted
contains silica
grains
Lecythis
grains
were
investigated.
and
oblong
sporadically
oblong
to 25
to
the ray
in
the ray
Grains of
ones
but there
cells,
no
silica
to the few
grains
were
probably
No
Grias
silica
(I),
grains
and
to
the
x
(Plate 1/3)
large,
size
of the
20/um.
The
grains
exceptions: Lecythis peruviana
parenchyma cells. Besides,
species
these
of Lecythis and their occurrence
in the other
species.
In L.
gigantea (two
observed. Dr. G.T. Prance (New York Botanical Garden)
studied the herbarium vouchers of these two
referable
occurrence
20,umto70
X
are two
cells and also in
grains present
sporadic
globular, always
from 45
much smaller than those in the other
abundant, contrary
samples)
or
species.
In his
opinion
the
samples
are
Eschweilera.
were
Gustavia
observed
in
Asteranthos (1),
Bertholletia (1),
Couroupita (8),
(4).
MELIACEAE
Silica
grains
species
gated,
other
occur
with and
in
two
species
genera of this
without silica.
six of these contained silica
shapes
are
family. Both Guarea and Trichilia comprise
A total of fifteen
grains
size of the
present too,
in the ray
species of Guarea
cells. Grains
globular grains
up
to
18 p
was
investi-
mostly globular but
m
and the size of the
115
B.
J.
H,
TER
oblong
ones
shapes
are
in the
grains
laurentii
grains
G.
grains
are
As in
There is
The
in
origin
species
of the
were
these
than
one
and
per
studied and
contain silica
Amos
to
grains.
grains
Nothing
T.
built
is
nineteen
grains normally
more
do
is
and
not
Amos
non-siliceous.
by
in the
to
same
than in Guarea.
in each cell.
not
known.
African
species,
(1952)
An
(up
is
contains silica grains in the ray cells.
restricted to the
&
Pennington
said
frequent
The
elegans.
occur
not occur
investigated by
as
T.
except in T. trinitensis
are
G.
species of Trichilia
and
grandifolia,
grains
investigation
(1952)
out of
12
to
and
reported
were
duckei, G. kunthiana,
G.
davisii,
grains
cell,
of Trichilia
specimens
According
loosely
like G.
species
,um). Species
casarettii,
built
loosely
grain
the distribution of silica
Trichilia.
genera
Trichilia,
more
compact
African
G.
three
Only
an
small silica inclusions
and British Honduras.
of Guarea without silica
15
to
In
large.
are
and up
globular
examined during the present
Apparently
of
both
me
costata,
T.
grains,
(1952)
Nigeria, Ivory Coast,
(up
G.
rhabdotocarpa.
without silica
Guarea,
never
lancei,
borisii,
for the greater part
are
jum).
are
G.
and G.
found
compact while those with various
are
to Amos
According
samples investigated by
pubiflora,
sample, but,
T.
globular grains
built.
loosely
alborosea,
G.
investigated
Two
the
15/i m;
x
less
again large grains
are
pohlii,
20
35
to
up
more or
in the ray elements of Guarea from
occur
The
WELLE
neotropical species
Styles (1975)
a
few other
these authors about their taxonomic
importance.
No silica
grains
were
observed in Cabralea (2), Carapa (2), Cedrela (2), and Swietenia
(3).
MENISPERMACEAE
The wood
anatomy
of this
in two
species
present only
the
contrary
three other
granular surface,
Silica
grains
size
up
were not
family
of
species
to
was
studied
Anomospermum:
of
this
13,um. They
by Mennega(1977).
genus
occur
Silica
proved
to
be
A. bolivianum and A. solimoesanum. On
lack
silica.
Grains
in both rays and axial
observed in thirteen other genera
globular
with
a
parenchyma.
investigated by Mennega.
OLACACEAE
The
occurrence
of silica
Liriosma. Grains
10 to 28
n
(1952)
this
quantities
No
and
116
m.
grains
globular
Some ray
and
in this
family
is restricted
sometimes oval
or
cells lack silica but contain
family is largely
to
a
crystal
non-siliceous, although
the ray cells of
with
oblong
a
instead.
some
one
genus,
granular surface,
According
species may
size
to Amos
contain small
of silica.
silica
grains
were
Ptychopetalum (1).
observed
in
Chaunochiton
(2),
Heisteria
(2), Minquartia (1),
SILICA
GRAINS
POLYGONACEAE
According
Parente
The
there
spaughia
investigation
with
they
8
grains
in
besides
(up
the
Triplaris
further research
so
No silica
grains
were
must
20
to
grains
of silica
in the
grains
ones,
m)
p
In
in
grains
besides
The
Triplaris.
and
shapes
they
are
with
a
prove if this is
only
one
cells
ray
in Neomill-
grains
are
are
slightly
sporadically
normally globular
smooth surface and
punctata where they
but
the
grain-containing
Ruprechtia they
other
globular
are
in T.
observed in Coccoloba
and,
8/um).
to
globular
large (up
small (up to 5 p m),
except
always
in
grains, especially
are
of silica
per cell
grain
Triplaris peruviana lacked silica grains,
available
is non-siliceous.
family
occurrence
occurrence
and small-sized
Symmeria
In
the
proved
10/i m). In this species,
The
are
this
(1952)
the
reported
more than one
cells without
granular surface.
a
m.
p
never
mostly globular
to
larger (up
also
are
are
present.
and Amos
(1950)
Triplaris gardneriana.
present
of four genera. There is
cells,
Chalk
for the first time
(1959-1961)
cells of
ray
Metcalfe &
to
sample of
measure
that
up
species
to
was
feature.
a constant
(13).
PROTEACEAE
In three genera,
cells
the
Euplassa, Panopsis,
and sometimes also
same
of the
genera
species
The
to
are
correct
10
the
/x
cells.
not
m,
In
(1952)
about silica
&
to
are
10 ,u
and
Engler
to one
Briggs (1975).
leoideae. An
&
two
13
/x
of
types
of the
m
ones
grains
voucher,
in
grains
nor
up
to
25/um,
var.
(1889)
all genera
occur
Euplassa
more
oval
same
are
as
or
less
like clusters of
,
Panopsis,
ones
and
montana
up to 38
25
x
that of those in the
simulans ( L.&L.2872) silica
however,
these results
did
(1)
shapes,
respectively. Only in Roupala
leaves
it should be referred to another
family
exception in
occur:
with various
cells is about the
On the
in
However,
distribution pattern, and size
in
some
doubt
as
its
to
species.
agreement
with
those of
contrary, Chattaway( 1948) reported nothing
Araujo
&
the genera
de Mattos Filho
containing
both classifications
in the rays and the
(1974a
comprising species
tribe, Grevilleae. Another classification
Here
grains
to
observed.
were
in the ray
correlation between these characters and the
of Panopsis rubescens
differences,
Prantl
Shape,
too.
The size
The herbarium
in this
grains
no
Mostly
in the ray
Mennega (1966).
to
which silica
occur
and 5
cells
parenchyma
surface; (2) grains
m
sample
minor
Roupala, species with silica grains
larger (globular
grains
one
present.
restricted
According
are
smooth
identification; possibly
Notwithstanding
Amos
5
grains
size of the
parenchyma
grains
a
genera.
particles glued together.
is 4
( O.N.S. 293)
(am).
or
with
globular, compact
small silica
axial
without silica
species
is very variable and there is
grains
delimitation of
Roupala
in the
and
was
proposed by
silica fall within the
is
one
and
1974b).
with silica
grains
Johnson
subfamily
Grevil-
sample of Petrophila teretifolia
pith, reported by Amos,
1952.
in
Nevertheless,
117
B.
J.
the
H.
TER
WELLE
of
occurrence
Proteaceae,
but it
No silica
grains
silica
grains has
be
to
seems
far been
so
important
an
in taxonomic
neglected
character in this
studies
on
family.
observed in Embothrium (1).
were
QUIINACEAE
The wood anatomy of this family
(1967). Silica
rence
grains
No silica
grains
be
to
were
extensively studied by
Gottwald& Parameswaran
reported for all species of Lacunaria investigated. Theiroccur-
were
considered
was
was
of taxonomic value.
observed in Froesia,
Quiina
and
Touroulia.
,
RHABDODENDRACEAE
Four
samples
of Rhabdodendron amazonicum
the ray cells and
a
to
30
grain
18 /um;
x
per
present
cell is
in the axial
probably
granular surface,
size of the
grains
were not
investigated.
cells. Grains
to
up
20
compact and others
are
sometimes
found,
too. The
parenchyma
globular grains
of them
some
were
two
grains
observed
per
cell,
u
Silica
deposits
are
not
present
oblong
ones
loosely built; normally
but ray cells without
by Prance( 1968,1972).
He
wood of Rhabdodendron differs from that of the Chrysobalanaceae
e.g. silica
occur
in
to oblong, with
globular
m, size of the
are
grains
even
in
up
one
grains
are
stated: ‘The
some
features,
in any form’.
RUBIACEAE
The wood anatomy of this
1969b,
1970,
1972,
family
was
studied extensively
by
Koek-Noorman
for the African genus Mitragyna
1974). Except
no
(1969a,
silica grains were
observed.
RUTACEAE
Only
are
two
species from
oblong
The
grains
more
than
No
one
silica
(1),
samples
are
size of 7
of
Galipea
globular, with
grain
grains
Cusparia (1),
carpus
a
to
20
Those in the axial
and less compact than those in the ray
per cell. Two
118
two genera contain silica grains. In Erythrochiton brasiliense
mostly globular with
were
(1),
cells. There is
investigated
and silica
granular surface (size
never more
occurs
only
in
one
they
oval or
than one
m) and
up to
are
grain
sample.
there is
never
per cell.
Esenbeckia
Rauia
a
were
parenchyma
and
observed
in Adiscanthus
(2), Fagara (1),
Ticorea
(1).
Helietta
(1), Balfourodendron
(1),
Hortia
(1),
(1),
Citrus
Metrodorea
(1),
(1),
Pilo-
SILICA
GRAINS
SABIACACEAE
Silica
grains
in
occur
genus not identified
silica
mostly globular
the globular
never
ones
more
In
samples
two
in the other two
sometimes
oval,
or
25 fim, size of the
to
up
than
of Meliosma sinuata and
samples
species.
particles glued together,
Grains
is
two
to
samples they
with
oblong,
show
a
normally
a
a
same
cluster of small
compact
structure.
size of
granular surface,
up to 50
of the ray
most
of the
samples
two
look like
grains
oval/oblong grains
per cell but
grain
one
the
x
20
cells
there
are
without
silica.
SAPINDACEAE
In this
family silica grains
neotropics, Toulicia,
means
that,
The grains
in the
species,
same
in
occur
the axial
contact with the vessels.
No silica
grains
seldom
occur
in which silica
was
samples
with and
show
various
There is
shapes
only
is
occurrence
one
genus from the
be found.
can
in the broad bands which
are
in
and their size is about 13 µm.
Allophyllus (2), Cupania (4), Matayba (3),
(1), Sapindus (1), Serjania (1), Talisia (10),
which
not constant
without grains
samples
especially
parenchyma,
They
observed in
were
(Amos, 1952).
observed. The
and Vouarana
Pseudima
(1).
SAPOTACEAE
One
or
from twenty-one
species
more
Manilkara, silica grains
were
The genus Chrysophyllum
other
neotropical
Gonggrijp (1932),
from Africa and
same
wonders
tropical
grain
Asia, species
see a
in
species
palaeotropics,
if, perhaps, taxonomists
are
grains
in Achrouteria
cell is often
species
and Sharma & Rao
without silica
have been
as
regards
more
Calocarpum
ones
show
various
a
size
difference in
shape
completely
shapes
of
up
grains
and size
Chrysophyllum species
grains
sporadic but they
are
In
one
genus
only,
occur,
to 30
x
are
to
with and
1
are
are
one
in the
genera from the
ones
globular
Normally
measure
up
all
neo-
and
one
neo-
there is
are
80
one
x
40/xm).
25 pm and the
are
parenchyma
globular.
without silica
to
to
Sometimes the grains
are
to
genera
normally present
always large (up
the
species
most
present.
grains.
S/xm). They
in
successful in the delimitationof
in the ray cells and the axial
(up
(1970)
their silica contents,
filled by the
18
without silica. In all
always present. According
genera than they have been with genera in the Old World.
Caramuri contains silica
no
with and
with and
grains
marked differencebetween all but
genera from the
The entire ray
In
silica
(1952), Murthy (1965)
Amos
investigated.
were
comprises both species
per cell but sometimes cells with two
The
oval
genera
observed.
investigated
genera
genus. We thus
tropics and
not
fragmented.
cells. There is
As mentionedbefore,
present.
No
grains
were
119
B.
J.
H.
TER
WELLE
observed in C. gonocarpum and C. viride. According
easily
are
separated
Grains
species.
globular,
the
schomburgkianum
30
upright
or
square
may
oval
15 to 23 /im, size of the
ones
be
upright
or
square
in the
well. Size of the
In E. balata the
17
x
Only
grains mostly
with
Amos
globular shape
a
globular,
of Lucuma
and
Gonggrijp (1932)
in the
10/xm) with
are
parenchyma
sample
one
10
to
25 pm, of the
cells
procumbent
are
ramiflora
and C.
marginatum
smaller than those in the
mostly globular
are
globular
shapes
but other
oblong
ones
up
to
smallerthan those in the
the
largest grains
found
are
cells.
ray
Globular grains(up to
Those in the axial
grains
are
species
containing,
size of the
In C.
/um.
ray cells
globular grains
grains
13
x
silica
other,
granular surface,
a
procumbent
J. C. Lindemanthese
from the
30
up to
ones
cells. On the contrary, in E.
ray
procumbent
tella the
oblong
in the
grains
with
oblong,
cells. In Ecclinusa the
ray
as
present
or
to Dr.
characters
morphological
on
smaller (up to 8
are
a
granular surface
but other
do
not
shapes
present
well. Size up to
as
differ from those in the ray
from Amazonas
ephedrantha
described this genus
(1952)
in Eremoluma. In Franche-
occur
are
frequently present. Those
than those in the square
investigated. Although
was
non-siliceous, silica grains
as
in the
ray cells
procumbent
upright ray cells which
or
lO/um.
cells.
measure
are
up to
15;um.
The
in
grains
to
10
measure
sporadically
show various
Micropholis
20 /xm.
Normally, they
then
present,
they
but
shapes
not
are
present
of them
most
in the
much smaller than those in the
are
globular,
are
procumbent
ray
and
if
cells, but,
upright
or
square
ray cells.
In Nemaluma
grains
are
In
the
15/xm,
up to
cells mostly do
globular grains
oblong
contain silica
not
seems
times
genera, the
be constant. There is
to
oblong
Twenty
cells,
are
lack
grains
two
20 pm,
occurence
species
In the
and P.
up to 28
x
the
shapes
are
15,um.
x
measuring
occur,
grains
one
Various
The
present.
The square
or
globular
upright
ray
in the axial
grain
per ray
shapes
are
or
up
to
15/xm. Contrary
of
parenchyma
parenchyma cell,
globular
present,
to
Neoxythece
but
grains
some-
up
to
15
were
to 25
procumbent
up to 28
are
of silica
normally
of Pouteria
pariry,
ones
shapes
grains.
occur.
mostly globular (18
grains.
mensalis,
per
ones
cell
of other
grains
various
Neopometia grains with
the other
and
pm).
ray
grains
The
cells
are
The
investigated.
marginal
of four
grains,
restricted
to
the
ray
ray cells in P. caimito sometimes
P.
species,
smaller than those in
glomerata,
the square
P.
krukovii,
or
upright
P.
ray
cells.
The surface of the
and
are
mostly globular.
granular surface,
oval
occur
Only
grains
in
one
and
measure
Radlkoferella.
sample
variation from
up
to
globular (up
up to
15/xm,
Various
shapes
The maximum size is about
of Richardiella
granular surface and
120
in Pradosia is sometimes smooth.
They
In Pseudocladia and Pseudolabatia they
to
20
was
yam)
15 pm
to
investigated.
oval
occur
(up
to
28
from
upto9/xm
globular,
globular
with
to more or
a
less
18 pm.
The
X
measure
are
shape
shows
a
continuous
15,um). Globular grains with
in Sandwithiodoxa. In Sarcaulus
a
grains with
SILICA
various
shapes
are
but globular
present
ones
frequently found.
most
are
GRAINS
Size up
to
15 /xm.
ca.
No silica
grains
observed in Manilkara (8).
were
SIMARUBACEAE
Silica
grains
shapes,
ones
occur
from
up
13
to
in
some
globular
/um,
oblong
smaller (max. 8 p m). The
of
species
oblong,
to
ones
are
up
grains
one
genus,
present.
18
to
10
X
were not
Simaba. In the
Size of the
grains
same
in S.
S. alata and S.
pm. In
observed
sample
guianensis they
Webber (1936) in her
by
various
multiflora: globular
study
on
are
the
wood anatomy of the Simarubaceae.
No
silica
grains
were
observed
in
Aeschrion
Picramnia
(1),
(3),
Quassia
and
(1),
Simarouba (1).
STYRACACEAE
Five
species of Styrax
Panama).
loosely built.
cells
The
more
than
Indonesia
in S.
grains
one
are
the
species,
shape
is variable.
grain
per
grains
They
occur
cell.
without silica.
sumatrana
and
investigated
although they sometimes
never
and
were
In the other
only
are
are
one
small
very
mostly found
sporadically in
According
In
is without silica (S.
to
the
small
(4
to 8
procumbent
Gonggrijp (1932)
from
and often
in the square
to Amos (1952) two
contrast,
argenteus
ray
or
upright
ray
cells. There is
species from Malaya
reported
very small
from Indonesia.
THEOPHRASTACEAE
All
are
a
samples investigated
from the genus Clavija contain silica grains in the ray cells.
mostly globular but other shapes
are
present
as
maximum size of 23 /u m, whereas they do not exceed
never
in C.
more
than one
grain
per
cell, but they
do
They
well. In C. lancifolia the grainsreach
in C. parviflora. There is
not occur
in all
ray
cells, particulary
not
parviflora.
TILIACEAE
In
one
neotropical
genus,
this genus, because L.
globular
with
of the ray
Luehea,
silica
grains
occur.
seemannii from Panama
to
smooth
surface,
size up
cells contain
crystals
instead of silica
a
7 /im,
Their
(Stern
et
presence
al.
never more
1839)
than
is
not constant
in
lacks silica. Grains
one
grain
per
cell, part
grains.
121
B.
J.
H.
TER
No silica
WELLE
grains
were
observed
in
Apeiba (3), Christiania (1), Heliocarpus (1),
Lue-
(3), and Mollia (2).
heopsis
THEACEAE
The
occurrence
of silica
Ternstroemia. Two
T.
circumcissilis,
mostly with
a
T.
No silica
delicatula,
T.
schomburgkiana,
5
max.
species lack
and T.
never
than
one
one
genus,
( T. browniana,
Grains
seemannii).
more
cells in
silica
globular
grain
per
cell,
and
but
contain silica.
not
grains
is restricted to the
ray
family
contain silica and 5 other
smooth surface, size
ray cells do
most
in this
grains
species
observed in Cleyera (1), Laplacea
were
and Pelliciera
(1),
(1).
VERBENACEAE
Silica
grains
lack silica
species
V.
censis,
cell.
V.
stahelii,
They
are
(
V.
cells in
V.
amazonica,
triflora).
and V.
V. compressa. Size of the
found in the
fibres of V.
septate
No silica
with silica
sample
grains
V.
grains
V.
krukovii,
grain present
size 20
/um).
up to 15 pm. In another
grains. Loosely
observed in
one
the contrary, 8 other
V. orino-
in every ray
One
sample
of
cells. Shape and distribution pattern is
contains many
sample
excelsa,
oblong (max.
in the
ray
megapotamica. Max.
were
of Vitex. On
species
V. cymosa,
In V. compressa there is
globular grains
silica is not
present. This
species
three
cooperi,
and sometimes
mostly globular
contains
floridula
like that in
in the
ray
occur
built
septate
grains
size 9
Their
sample
of the
fibres which were
of various
shapes
hardly
occur
in the
is abundant.
occurrence
Aegiphila (2), Citharexylum (3),
same
and Petraea
(1).
VOCHYSIACEAE
In this
Amos
family
the
grains
silica, Normand(1966,
vary
ones
up to 25
occur
and
shape
as
large
ingens
variable.
122
X
as
they
13
sporadically
ingens and Q.
var.
but
used
grains
are
In
10
rosea.
are
to
the
Q.
grains
Q. cordata,
near
are
are
family
most obvious in
acuminata
grains
cells of
one
genus,
Qualea. Although
may have small
quantities
of
thischaracterinhiskeytodeterminetheVochysiaceae
they
small
loosely
(3
globular
are most
the vessels in
a
to
5
and
Q. albiflora
Size of
Globular grains with
The
ray
this
species of
mostly globular.
in the rays
too. These
In
restricted
some
1967)
from the Guianas. The
in
are
reported that
(1952)
Q.
ones
frequent
Q.
acuminata.
up to
near
the vessels. Grains
coerulea. Grains of various
smooth
/xm).
surface
Another
built and show a
occur
type
in
is found in
were
shapes
Q. ingens
granular surface,
Q. cryptantha, and Q. dinizii silica grains
They
of oval
15
their
not
var.
Q. ingens
shape
is
observed.
abunda t
proba ly
taxa
grains
to
respctively:
resticed
abunda t pres nt, proba ly abunda t cells
of
-----
r
e
s
p
c
t
i
v
e
l
y
:
pres nt,parenchyma, pres nt,resp ctively: pres nt,
s
i
l
c
a
neotrpical
sil ca marginal
the
of
of
Distrbuon
the
grains ray
r
r
rr
r
Distrbuon
secondary
in
Locality
,
Amazonas
Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam Brazil
Br.
,
r
(p)
(r)
i.
rr
V
18
Colection
in
grains
Forestir
of
et
Table
1.
in
TABLE
Agricole
Guiana
Suriname
Jonker
Fr.
&
Departmen,
de
Maas Behe r
&
Bos
-
Horeus Lindeman Norde
&
Maas
81
5780
Hulster
BW de
Lindeman Lindeman Stey rmak
Hanco k
f.
Hook.
ex
's
-
-
-
-
-
-
-
-
Hgi anteum anco k Benth. Ducke sagoti
-
f
f
f
f
f
L.
H.
Jonk.
a.
Spruce
&
and
Abreviatons Colection BAFOG
10735
Stahel O.N.S. Stahel LBB Kjrukof Stahel
&
Bos
_
4723
89343
&
Lands Oldenburge,
B
o
s
c
h
w
e
z
n
F
l
o
r
s
c
h
ü
t
z
G
u
y
a
n
i
s
,
Lindeman Lanjouw Prance
D
a
n
i
e
l
s
B
u
r
e
a
u
F
o
r
e
s
t
B
r
a
z
i
l
locality
Behe r
278 590 268
6057 6194
Schulz
Haas
Guyana
;
used
rr rr
r,
P
,
the
Distrbuon
r
rr
b.
1:
rr
i
f,
(f)
sil ca
r
the
xylem
PP
rr
tyloses
fibres, pres nt,axial proba ly rays, pres nt, rays, the present ves els
ff
rr
Ms.
de
Ms.
&
H. L.
&
&
BBS Br.
BW
Dan. F.D. Flor.
&
LBB
&
L. L.
O.N.S. Pr.
Species
Hook. Hook. Hook. Hook. Hook.
Anacrdiea Anacrdium gianteumocidental spruceanmtenuifolm Loxpterygium sagoti sagoti sagoti sagoti sagoti
123
B.
J.
H.
TER
WELLE
v
Distrbuon
rr
(r)
r
r
rr,
,
rr
r
r
rr
r
rr
r
t
v
p,
pp,
P,
/
pp pp
Locality
r
rr
p
pp pp pp pp pp
pp, PP/ pp, (p)
rr rr rr rr
Amazonas
Amapa
Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam Brazil
Br.
Br.
r
r
r
rr
rr, ff, ff,
Amazonas Amazonas Amazonas
Guyana Guyana Guyana Guyana
Br. Br. Br.
rr (f> ff
f,
p,
P/ p,
(p) p, pp,
P p
ff, ff, ff/ ff, ff, ff, ff, ff, ff,
Amazonas
Brazil
pp,
inam
Surinam Surinam Surinam Surinam Surinam Surinam Surinam Sur Surinam Surinam Surinam Surinam Surinam Surinam Guyana
Br.
869
301
Colection
5184 8927 2420 2761 3568 1889 6345 5721 3241 6288 48453
Stahel Maguire Schulz
L. L.
&
&
L. L.
L.
L.
&
&
L.
L.
Lindeman Lindeman Krukof Lindeman Maguire
17628 45 21 46107
5143
158 306 257
F.D.
Maguire Maguire Maguire Ducke Ducke Ducke
9182
Pires
308
7142
Stahel Krukof
4671 2410
Lindeman
12
180
L.
&
L.
23
390
Hulster Hulster
L.
Stahel
&
de L. de
(Aubl.)
(contiued)
aquaticum
1
Table
Species
124
Vischer
Schum.
(Aubl.)
Cuatr.
Uit . Uit . Uit . Uit .
Ulbrich Ulbrich
Huber
duckei
Benth.
Aubl. Aubl. (Schum.)
multifora
Benth.
Benth. Benth. Benth. leiantha Benth.
belmn si
Ducke
sp.
cf.
&
Schulz Stahel Dan. Lindeman Schulz Schulz
L.
2139
&
L.
F.D.
robust m robust m Engl. Engl.
W.
March March
(Swart) Swart
firmum
6778
Jonk. 8332 8937 693
Swart Swart
Rodr.
Schum.
8330 262
apiculatm
a
q
u
a
t
i
c
u
m
crasum nervosum nervosum nervosum spectabile spectabile guiane si guiane si lasiocalyx Bonetiac multifora multifora multifora Haploc thra paniculat verticla
Bombace Bombax
Quar ibea
Kielmyera Burseac Dacryodes Parpotium Protiura
Archytea
var. var.
(Aubl.) (Aubl.)
Engl.
Engl.
Swart Swart Swart Swart
(Turcz.) (Turcz.) March March March
heptahylumheptahylum insigneneglctum neglctumneglctumneglctumpolybtrumpolybtrum sagotianumsagotianumsagotianum
SILICA
rr
GRAINS
t
v
r
op,
r
f,
rr
ff,
rr
r
'
f
rr£
,
(f) Pf
P.r
rr,
rr
r
p,
,
rr
(p) p,
r
Amazonas
Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam
Br.
Surinam Guyana
rr rr
rr rr
rr
rr rr rr rr
Amazonas
Amazonas
Br.
Br.
Amazonas
Surinam Surinam Surinam Surinam Surinam Surinam Surinam
3077
2377 3065
rr rr rr rr rr rr rr rr rr rr
Amazonas
Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam
Br.
3794
3753
Br.
238
:
3954 4525 6117
8341
40
1041 1042
Stahel Lindeman Lindeman Lindeman BBS BBS Schulz
1719 6168
L.
&
L.
7017
8304 5227
24576
2991
Lindeman
Krukof Maguire F.D.
6795 6857
Donselar Donselar Donselar Donselar Donselar
Krukof Lindeman van
van
Krukof Lindeman van
van
van
7356 6851
2561
L.
Schulz Krukof
&
596
4909 1573 2694 6163 9014
Heyligers Stahel/GongL.indeman Lindeman Krukof
L. L.
&
L.
&
L. L.
(Radik.) Foreo (Radik.) Foreo
spadlcea spadicea
Smith
rhoif la rhoif la
Mart.
burseifola
&
Gleason
var.
Meyer
Benth.
ssp. ssp.
Mart. Mart. Mart.
Sandw. Sandw. Sandw. Wild. Wild.
amzonicus Gily G.FW.
Benth.
cuspidat
sp.
b
u
r
s
e
i
f
o
l
a
m
a
z
r
u
n
e
s
i
b
u
r
s
e
i
f
o
l
a
b
u
r
s
e
i
f
o
l
a
t
r
i
f
o
l
a
t
u
s
h
o
i
f
l
a
r
h
o
i
f
l
a
d
e
m
r
a
e
d
e
m
a
r
e
d
e
m
a
r
e
C
a
r
y
o
c
e
a
Burseac Tratincka
Anthodiscu Conarce Pseudocnars
cf.
sp.
Rourea
var.
Radik, Radik, Smith
(DC.) (DC.) A.C.
cuspidat
cf.
Peyr.
Miq. Miq.
pubesc ns pubesc ns rectinrva surinames surinames
Hil. Hil.
amzonicum Peyr.
St. St.
Benth.
Cav. Cav.
Bongard
Mart. Peyr.
Erythoxlace Erythoxyln amzonicum citrfolum citrfolum macrophylum acrophylum icranthum nitdum parense
amplum
125
B.
J.
H.
TER
WELLE
Distrbuon
rr rr rr rr
r
rr rr
r
(r) rr
rr
rr rr rr rr rr
rr rr rr
(r)
rr rr rr rr rr rr rr rr rr rr rr rr rr rr rr rr rr rr rr
Cat rina
Locality
Amazonas Sta. Amazonas
Br. Br. Br.
Colection
5551
2 181 10 61
Ms.
Pr.
Amazonas Amazonas Amazonas Amazonas
Br.
Br.
Surinam Surinam Surinam Surinam Surinam Peru Brazil Brazil
139 2
&
Amapa
Reitz Krukof Schulz
27
7365
51745 1869 6997
Stahel Schulz Maguire
L.
&
L.
24279 5192
8193
Br. Br.
Br.
6922 7171
7126
8050
Lindeman MADw Pires Krukof Krukof Krukof
USw
USw
Surinam Surinam Surinam Guyana
Amazonas Amapa
Br. Br.
Surinam Surinam Surinam Guyana Surinam Surinam Guyana
57527 5 864 45903 6316 51724 2856 2481
35636
Yw
Maguire Maguire Maguire Krukof Maguire
L. L.
&
427
10721 5540
&
L. L.
L.
&
LBB
F.D.
L.
165
3684
Amazonas Amazonas Amazonas Amazonas
Amazonas
Br. Br. Br.
Br.
Surinam Surinam Surinam Brazil Brazil
Br.
6384 7205 8714 3840 8975 4149 8958 47079 6375 471 1
Stahel F.D. Krukof Krukof Krukof Lindeman Krukof Lindeman Schulz MaguireKrukof Maguire
Arg.
Hof m.
et
,
Pax
Muel. Arg. Arg.
Arg.
Muel.
amzonicus (Spreng.)
Pax
Saldnha (Klotzsch)
Aubl.
(Didrchs.)
Muel. Muel.
Hof m.
Walp.
Tr. Tr.
et
Pax
Benth. Ducke Ducke Ducke
)
(Mart. Walp.
Rich.
(Mart.) Engl. Sandw, Vesque Vesque
Rich. Rich.
L.C.
aff.
et et
Tr. Tr.
Smith
Tr.
et et
PI. PI.
Tr.
vel.
Walp.
k
a
r
s
t
e
n
i
a
Ducke Benth.
g
u
i
a
n
e
s
i
b
r
a
s
i
l
e
n
Poep. Poep.
(
D
i
d
r
c
h
s
.
)
(
D
i
d
r
c
h
s
.
)
C
r
o
i
z
a
t
e
l
a
t
a
Sandw. Sandw.
(contiued)
palmicda
b
r
e
v
i
s
t
a
m
n
c
e
p
h
a
l
o
s
t
i
g
m
a
c
e
p
h
a
l
o
s
t
i
g
m
a
s
i
p
h
o
n
i
d
e
s
b
r
a
s
i
l
e
n
s
c
h
o
m
b
u
r
g
k
i
s
c
h
o
m
b
u
r
g
k
i
s
c
h
o
m
b
u
r
g
k
i
m
a
c
r
o
p
h
y
l
a
m
c
r
o
p
h
y
l
a
m
a
c
r
o
p
h
y
l
a
l
a
n
c
e
o
t
u
s
m
c
r
o
p
h
y
l
a
c
a
l
o
d
i
c
t
y
o
s
g
u
i
a
n
e
s
i
g
u
i
a
n
e
s
i
p
a
l
m
i
c
d
a
p
a
l
m
i
c
d
a
p
y
r
i
f
o
l
i
a
c
o
n
l
o
r
c
a
r
i
n
a
t
k
r
u
o
v
i
secunda secunda speciosum stigmatos umbelat
n
i
t
d
a
e
l
a
t
a
e
l
a
t
a
gr
a
t
a
gr
a
t
a
E
u
p
h
o
r
b
i
a
c
e
A
c
t
i
n
o
s
e
m
n
S
e
n
f
l
d
e
r
a
O
e
d
m
a
t
o
p
u
s
Gutifera Clusia Tovmita
Maprounea Micrand
Table Species
1
126
Aubl. Aubl.
L.C. L.C.
sp.
Eyma
et
PI.
PI. PI.
ap. ap.
et
PI.
SILICA
rr
rr rr
p, p,
(P) pp,
pp, pp,
Surinam Surinam Surinam Surinam
Fr.
rr
Br.
guianesi
Hatschba
L.
L.
Krukof
Benth.
ff ff ff
rr
Amapa
r
51845 Jonk. Jonk. 5 307
2704
&
p,
t
rr
,
r
(p) rr rr p,
v
rr,
Amazonas Amazonas Par na Amazonas Amazonas Amazonas Amazonas Amazonas
Colmbia
Br. Br. Br. Br. Br. Br. Br. Br.
Br.
1638
320
56549 5221 5317
H.
24548 2282 5930
51738 7193 5597 de 5568 8796 8164 8690
L.
H.
3871
&
Maguire Maguire Stahel MaguireKrukof Krukof IANw Maguire Lindeman
&
F.D. Dan. Dan.
L.
9020
Maguire Krukof Krukof Krukof Krukof Krukof Krukof USw
209 9
&
L.
IAN
Cuatr.
guianesi guianesiguanesi guianesih pidula
var.
rr rr rr rr rr
(p),
804 866
13878
18
BAFOG Lindeman Lindeman Stahel
rr rr rr rr rr
Amazonas Amazonas Amazonas
Par na Amapa
GuyanaBr. Surinam Surinam Surinam Surinam Brazil Br. Br. Br. Surinam Surinam Surinam
Br.
6252 6381 2869 6653
&
r
rr
Amazonas
Guiana
1239
rr
rr rr rr
.
GRAINS
Taub.
Mez
var. var. var. var. var.
ascheronia
guianesi
Benth. Benth. Benth. Benth. Benth.
g
u
i
a
n
e
s
i
g
u
a
n
e
s
i
g
u
i
a
n
e
s
i
g
u
i
a
n
e
s
i
g
u
i
a
n
e
s
i
Humirace Sacogltis
Lauroea
Allen Allen Allen Allen
C.K.
Kostera. C.K. C.K. C.K.
maguirean
(Samp.)
Cryptocary Licari
ra huba
wilhemnsi wlhemins wilhemnsi
Mez
(Meis n.) Taub. Taub. Taub. Kosterm.
itauba (Meis n.) (Meis n.) (Meis n.) (Alen)
Mezilaurs
itauba itauba itauba syna dra
(Meis n.)
Miers
Mez Mez
(Meis n.) (Meis n.)
glaucina
Ocotea
splendens plendens Lecythidae
(Beg.) Ducke
lineat decandra
Alantoma Carin a
O.K. O.K.
Miers
(Mart.)
(Rad i) (Rad i)
Smith
Rodr.
W.
Ducke Ducke A.C. Miers
domestica estrl nis estrl nis micranth multifora pachyanthpyrifomis
rimosa
Crytophra
127
B.
J.
H.
TER
WELLE
v
Distrbuon
t
v
t
t
C
v
v
r
r
r
r
r
rr r, rr rr
Amazonas Amazonas
r
r
rr rr rr
Rica
Locality Surinam Surinam Surinam Surinam Surinam Costa Venzuela Surinam Surinam Guyana
Br. Br.
Amazonas Amazonas
Br. Br.
rr,
rr
rr
rr,
rr
rr
.
p, rr
pp,
rr p,
pp,
rr, rr
,
rr
,
<
6220
Colection
1653 1513 2198 7399
49
44
Lindeman Stahel Krukof Krukof Stahel
L.
&
L.
1561
Allen Bruijn
L.
Schulz P.H. Stahel
de
1163
a
2483
7254 8893
Smith
&
A.C.
L.
Krukof Krukof
Knuth
Sandw.
gloriosa
(conti ued)
1
Table
Species
Lecythidae Courati
128
Smith
Smith
Aubl. A.C.
A.C.
B
2564 1077
517 1
7687 8338
F.D. BBS Schulz
Standi. Standi.
6450
10898 4806
8327 2308 Jonk.
30
1022
Maguire Schulz F.D. Krukof Schulz
LBB
Smith
Eyma Eyma ex
(Smith) (Smith) Ducke
Amazonas Amazonas Amazonas
Guyana Surinam Surinam Br. Surinam Surinam Guyana Br. Surinam Surinam Surinam Surinam Surinam Guyana Guyana Ven zuela Br. Br. Br. Br. Surinam Surinam Surinam Br. Br. Br.
3561
137
rr
,
Amazonas Amapa Amazonas Amazonas
30147
USw
rr rr rr
(p) p, p,
rr (P) (p) (p) rr rr
rr p, P/ p, rr rr (p) rr p, rr
Amazonas
Amapa
rr
rr rr
BBS LBB
L.
&
L.
2518
40502 3870 8399 51861 4847 6233
7662
A.C. Ndz. Ndz.
A.C. A.C. A.C. A.C. A.C.
alata
(Aubl.) (Aubl.)
Miers
A.C.
Mart.
Eyma Eyma lora
2164 6299 7183 4848
L,
Bret ler
Dan. F.D. Maguire Krukof MaguireKrukof Krukof S«hulz Stahel Krukof Krukof Krukof
&
&
L.
Smith
Smith Miers Miers Miers
Smith Smith Smith Smith Smith
a
45
(Poit.) (Poit.) (Poit.) Sandw. Sandw.
Knuth
Ducke Smith Smith
Sandw. Sandw. Sandw. (Huber)
A.C. A.C.
Miers Miers A.C.
Eyma
Miers Miers
(Poit.) (Poit.) (Miers) (Berg) (Berg)
guianeskiru ovi macrospema ultifora multifora obl ngifolapanmensi panmensi stelat stelat stelat stelat stelat Eschweilra amar amar blancheti a colina colina conferti coriace corugat corugat corugat decolrans decolrans grata grata grata iqutoseni arana krukovi kru ovi labriculat longipes longipes obtecta obversa obversa
j
SILICA
GRAINS
t
v
rr,
rr
rr
rr rr rr rr rr
Surinam Surinam Surinam
a
136
5193
(p),
Amazonas Amazonas Guiana
Br. Br, Fr,
r
r
r
rr
(p) rr
p, rr rr P.
r
r
r
Amazonas Amazonas Amazonas
Pan ma Surinam Surinam Surinam Surinam Ven zuela Br. Br. Surinam
3770
8352 1665 7116 1049
Br.
Surinam Guyana
r
r
r
r
Peru Br.
Surinam Surinam Surinam Surinam
1130
3889 4585 6306
2 10
Krukof Lindeman Maguire BAFOG Krukof Krukof Maguire MADw Krukof
Ducke
Mart. Men ega
Miers Miers Miers Miers
(Spruce) (Sagot)
Ben.
Ben.) Ben.)
(Steud.) (Steud.) (Steud.) Smith Smith
Knuth (R. (R.
A.C. A.C.
(Huber)
R.
jar num
245
Guiana Guiana
Fr. Fr.
PP. PP.
223 260
Smith Smith Huber
Wms.
rr
.
M M
36
Stahel Lindeman Lindeman Lindeman BAFOG BAFOG Stahel BBS
60
(Aubl.)
Sandw.
,
(p) (p) rr rr rr rr rr P. rr rr rr rr
P.
Guiana
Guiana Guiana
Fr,
Fr. Fr.
Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam Guyana
Steud.
Miers Miers Miers
Eyma Eyma
pp pp pp pp pp pp
Fr. Br. Br. Br.
M
Stahel Lindeman Schulz Krukof Krukof BAFOG USw BBSLindeman BBS Schulz Bret ler Krukof Kurkof Lindeman
r
Amazonas
5097 1531 6954 1995 3703 405 1 4630 4811 51864
8335 7108
104
r
Guiana Amazonas Amazonas Amapa
158
698 170
p,
rr rr
rr£ rr
rr
,
389
A A
L.
1070 1071
&
L.
BBS BBS
5840
Venzuela
8 349
M
9462
Lindeman Schulz
N
1 030
LBB
89
8238
28
3311 252
1197
BAFOG Stahel Schulz F.D. BAFOG BAFOG
Steyrmak
Ben.
Steud. Steud. Steud. Steud. Steud.
Amshof
R.
lorum
(Aubl.) (Aubl.) (Aubl.) (Aubl.) (Aubl.) Amshof Amshof albif
Ben. Ben. Ben. Ben. Ben.
Benth. Benth. Benth. Benth. Benth. Benth.
g
u
i
a
n
e
s
i
guiane s
wachen im davis
pachysepal subglanduosa ubglanduosa ubglanduosa truncat truncat Holpyxidum davis davis hians hians parensi peruvian usita Leguminosae guianes guianes guianes guianes guianes guianesi guianesi Sclerobium albiforum albiforum albiforum albiforum albiforum guianes guianes guianes guianes guianes guianes
odora odora odora odora persi tens pit eri simorum slmiorum
Dlcorynia
Lecythis
Dialum
(Poep .) (Poep .) (Poep .) (Poep .)
Sandw. Sandw.
A.C. A.C.
L.
Miers
R. R. R. R. R.
of.
129
B.
J.
H.
TER
WELLE
Distrbuion
rr rr rr rr rr rr rr rr rr rr
Locality
rr rr rr rr rr rr rr rr rr rr rr rr rr rr rr rr rr rr
Amazonas Amazonas Amazonas Amazonas Par na Amazonas Amazonas
Amapa
Brazil Br. Surinam Surinam Surinam Surinam Surinam Surinam Br. Br.
Br. Br. Br. Br. Br.
r
rr rr rr rr
Amazonas Amazonas
Amazonas
Br. Br.
Br.
Surinam Surinam Guyana Surinam Brazil Surinam Surinam Surinam Guyana Surinam Surinam Guyana Surinam Surinam Ven zuela
639
H.
51840
Colection
3866
IANw
Maguire
1070
1 025 10717
BBS LBB LBB
1159
1254 1464 1528 6492 4631 4716
L, L.
O.N.S.
&
&
L. L.
de
5409 1917
H.
365
480
3595
1021 6866 8314
47624
297
1 02 1 046 4023
182 7308
s.n.
5208
432
3536 4711
Krukof Krukof Krukof Krukof Krukof Krukof Stahel Men ega F.D. Krukof Krukof Schulz Maguire Stahel LBB F.D. Stahel Schulz F.D. Geyskes O.N.S. Bret ler Krukof
&
LBB
L.
DC.
DC.
C.
Rusby Rusby Rusby
Ducke
(contiued)
1
Table
carinat
Species Meliac e Guarea
130
DC. DC.
DC.
C.
C. C.
Pul e Pul e
ex
Wils. Wils.
(Jacq.) (Jacq.)(Briton) (Briton)
gned si gom a gom a grandifola grandifola guara guarrausbyi rusbyi
cardenasi
Trichlia
DC.
Rusby Jus .
Smith
Harms A.C.
Radik. Radik. Klotzsch
DC.
C.
Smith Smith
C.
DC. DC.
A.C. A.C. C. C.
(Miq.)
DC. DC. DC.
C.
(Miq.) Jus . Jus . Jus . DC.
A. A. A.
C. C. C.
C.
Rusby
cardenasi catigua ernesti froesi fusce ns fusce ns guiane skiruovi krukovi moritz moritz pro inquaro iman roaimna roaimna sube ilfoa surinames trinesi trinesl trinesi verucosa virdis
A.
SILICA
rr rr rr rr
rr
Amazonas Amazonas Amazonas
Br. Br.
Surinam
Br.
unk own
rr rr rr rr
rr rr
Par na Par na
Amazonas Amazonas Amazonas
Br. Br.
951
Krukof Krukof O.N.S. Krukof
1 595
de de
MADw
&
rr
r
Venzuela Panma Ecuador Panma
r
rr rr
Amazonas Amazonas Amazonas
Br. Br. Br.
r
rr rr
Surinam Surinam Surinam
1 962
2002 5143 1412 6749 5456 3653
H. H. L.
&
Br. Br. Br.
r
1716
H. H.
6328 6910 1285 6205
Surinam Venzuela
rr rr
GRAINS
&
L. L. L.
6249 8423 5277
1859
Bret ler Krukof Krukof Krukof Bret ler Stern Acosta-Sli Krukof Krukof Krukof Stahel
50
103
1053 1054
USw
BBS BBS
Steyrmak
Blk.
&
)
(Gros .
Spruce
adherns A.C.
Olac ea Lirosma
emarginat
Smith
Engl.
Miers
cerif a guiane sipalida
Stand.
Meis n
Meyer Benth.
Briton
Eyma
laxiflora Meisn. jnensi(Jacq.) paniculat Benth.
bolvian
Mey.
&
Wed .
Cham. Fisch.
(Bertol.)
Cham. Cham. Cham.
Meis n. Meis n. Stand.
NPolygnace eomilspaugh Ruprechtia laxiflora ramiflora Symeria paniculat- Triplaris carsna cuminga guayqilensmaenodr pavonipavoni puncta surinaes surinames surinames
marowi
131
B.
J.
Distrbuon
H.
TER
WELLE
rr
rr
rr
rr
p,
rr rr
rr rr rr rr rr
,
(p),
p,
r
rr (p) rr
p, p, p, p, p,
Cat rina
Locality
,
(p), (?) rr
(p)
rr
P/
rr
r
rr
r
r
p
p
p
p
<p)
Cat rina Cat rina
Sta.
Amazonas Amazonas
Br.
Br.
Br.
Surinam Guyana Surinam
Guiana Par na Sta. Sta.
Fr. Br. Br. Br.
6370
1934 7236
&
3040
1322
Krukof Krukof Stahel F.D. Lindeman BAFOG
Reitz
de
Surinam Surinam
6030
Klein
249 293
L.
Reitz Reitz Stahel O.N.S.
Peru
2497
10694
Surinam
3437
Par na Par na
Br. Br.
341
4923
Surinam Surinam
928
Surinam Surinam
Amazonas Amazonas Amazonas
Br. Br. Br.
1164
H. H.
47524
Schulz LBB Fanshawe Irwin
Wilams Donsela r
de de
H. H.
&
L.
van
Jonk. Jonk.
&
&
L. L.
368
Dan.
Dan.S
4687 4914 5330
10732
Stahel LBB Krukof Krukof Krukof
Huber
simulans Macbride
var.
Br.
6761
8322
&
H.
&
Surinam Surinam Guyana
a
H.
6413
291
Amazonas
28181
2827
Klein
Colection
rr
-
Benth.)
simulans Macbride
ex
Huber Huber Huber
Mart.
&
Ducke
Sieum.
cantreia rubescn
(contiued)
1
Table
(Schot)
Species Proteac Euplas a Panopsi
132
var.
Ducke
(Schot)
rubescn
(Spruce Benth.) Benth.) Benth.)
Sandw. Sandw. Sandw. Sandw.
Pohl.
ex ex ex
amzonicum (Spruce (Spruce (Spruce brasilen
(Rich.) (Rich.) (Rich.) (Rich.) Sieum.
Pohl.
brasilen
Nees
Aubl. Aubl.
Aubl.
trifolat
Radik.
Urb.
Radik. Radik. Radik. Radik.
sinuat Urb.
se ilfoa se ilfoa se ilfoa se ilfoa catr um acrophyla monta monta Rhabdoenrc Rhabdoenr amzonicum amzonicum amzonicum Erythocin sinuat
Rutace Galipea Sabicea Meliosma
Roupal
sp. sp.
pulvinat
Sapindcea pülvinat reticulat reticulat reticulat
Toulica
SILICA
rr
rr
r
r
rr
p,
rr
rr rr rr rr rr rr
rr rr rr rr rr rr
Amazonas
Amazonas
Par na Par na
rr
p,
Guiana
Guyana Surinam Pan ma Brazil Br. Surinam Br. Br. Guyana Surinam Br. Surinam Surinam Fr. Surinam Surinam Surinam Surinam
rr
rr
»
(p)
rr
rr rr p, rr rr
Amazonas Amazonas
Br. Br.
rr rr
rr rr
Guiana
Guiana
Fr.
Fr.
p,
,
GRAINS
rr
rr
r
(p) P,
(p)
Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam Surinam
1347 3312
51796
2506
913
L.
696
&
F.D.
L.
USw
H. H.
5593 54831 de de
H. H.
Maguire Krukof Maguire
Eyma
pomifera
Aubr.
&
L. L.
(Ducke)
186
L. L.
Maguire Krukof Stahel Stahel BAFOG
&
&
L. L.
L.
&
L.
Smith
Radik, Radik,
A.) A.)
acrenum
et et
233
5422 7752 24739
14
1304
4837
2160
a
M
134
13
55
L.
198
10802
7327
30
Stahel Krukof Krukof Maguire Stahel BAFOG Stahel BBS Lindeman BAFOG Stahel LBB Schulz Stahel
&
L.
Monach
Aubrév. Aubrév.
DC.
Mey. A.
Ducke
tomentosa Engl.
var.
(Rudge) Eyma DC.)
(Piere)
Pel gr.
Aubrév.
var. var.
A.C.
(L.)
F.D.
177 91
2221 463 2651
(Miq.)
et
Pier e
4757
M
marginatum arginatum
Pel gr.
Eyma
&
24309 7208
(Eyma) Smith
Bail .
gongrijp!
A.C.
Pier e
Pier e
DC.)
Pier e Eichl.)
(A.
Pier e
DC.)
et
et
Eyma
Pel gr. Aubrév. Aubrév. Aubrév.
Pel gr. Pel gr. longifola
Aubrév.
et
et
(Eyma) Aubrév.
(Eyma) Aubrév.
(Sandw.) Eichl.)
et et
Aubrév. Aubrév.
var.
et
(Eyma)
p
t
y
c
h
a
n
d
r
a
cladntha (Mart.) Pel gr.
(
E
y
m
a
)
G.FW. bal ta
sagotian ephdranth
engleri
(Eyma) (Eyma)
s
c
h
o
m
b
u
r
g
k
i
a
n
m
s
a
n
g
u
l
o
e
n
t
a
e
u
g
n
i
f
o
l
a
m
a
r
g
i
n
a
t
u
m
a
r
g
i
n
a
t
u
m
c
u
n
e
i
f
o
l
a
g
u
i
a
n
e
s
i
guianesimartina venulosa engleri ptychandra robusta
a
m
i
f
l
o
r
a
p
r
i
e
u
r
i
a
u
r
a
t
u
m
n
l
t
i
d
u
m
C
h
r
y
s
o
p
l
u
m
F
r
a
n
c
h
e
t
l
a
M
i
c
r
o
p
h
l
i
s
Sapotcea Achrouteia Calocrpum Carmuri
Eclinusa
Eremolu a Luc ma
Nemaluma Neop metia Neoxythec
pomifera ma osum
Achrouteia op oslta
vs.
(H. (H.
Miq.
Mart.
(A.
egensi Pier e (Mart,
(A.
dura dura
133
B.
J.
H.
TER
WELLE
Distrbuon
rr
rr rr rr rr rr rr rr rr rr rr rr rr rr
rr rr rr rr rr rr rr
rr
Amazonas Sta. Amazonas Amazonas Amazonas Rica Amazonas Amazonas
Amazonas Par na Guiana Amazonas Amazonas Rica
Amapa
Br.
Br. Br.
Costa
Br.
r
r
rr rr
rr
rr rr rr rr rr
rr
rr
rr Pi
Cat rina
Locality
Surinam
Br. Br. Br.
Surinam Costa Surinam Surinam Surinam Surinam Surinam
Br.
Br. Br.
27498
Guyana Surinam Surinam Guyana Surinam Surinam Surinam Surinam Guyana Br. Surinam
s.n.
1322
9
&
L.
Allen
1505 5700 7453 24310 24596 5034 de 671 6344 5283
M
Krukof Reitz Krukof Krukof Stahel Krukof P.H. O.N.S. Krukof Krukof Schulz Maguire MaguireKrukof
L.
H. L.
&
&
L. L.
517 4
Lind. 2408
Allen
937
BAFOG Krukof Krukof P.H.
MaguireF.D.
2
&
Br. Br.
s.n.
H.
434
Amapa Amazonas
39094
3494
5124 670 Klein 5177 6632
Colection
Fr. Br. Br.
Amazonas
&
L.
220
3759
95
Cowan
204
&
L.
F.D. BBS BBS
7293
1041
122
4846
51839 1419
Stahel Schulz F.D. Krukof Stahel MaguireKrukof
Huber
Baehni
glabrescn
Smith) Radik.
var.
(A.C. Pav.) Radik.
Baehni
Aubrév.
et
Pier e
et
Wms.
Cronquist (Briton)
Wms.
Baehni Baehni
Radik.
L.
&
Radik,
Pel gr,
L.
Baehni
Smith) Smith)
(Spreng.) Eyma Eyma Baehni CronquistStand. Ducke
DC.)
Aubrév.
Aubrév.
(Eyma)
Aubrév. Pel gr.
et
Aubrév.
Pel gr.
f.)
Aubrév. Aubrév. Aubrév. (Sandw.)
(Gaertn.
Eyma
Radik.
minutflora (Eyma) filpes brachyndra Aubrév. (Eyma) (Eyma) (Eyma) rivcoa egregia brasilen (Mart.)
prealta
(contiued) anibefolia
guiane si guttahetrodxa hispida inflexa kruovi melanopda mensalis pari y salicfoa surinamesi urinames trichop da trilocuaris tripla foia schomburgkian Pseudoclai scytalophra Pseudolabti Radlkofer a brachyndra trigonsperma tigonsperma tigonsperma Richardel Sandwithoxa macrophylus
c
a
s
i
o
c
a
r
p
g
l
o
m
e
r
a
t
c
a
i
m
t
o
e
x
c
e
l
s
a
Sapotcea
et
Smith)
Baehni
&
(Mart.) (Miq.) Aubl. Stand.
Eyma
(A.C. Baehni
(Ruiz, (A.C.
Eyma (A.C.
(Ducke)
(Ducke)
Baehni
1
Table
Species Pouteria
134
nuda
Pradosia
(A.
Sarculs
SILICA
rr rr rr rr rr
r
rr rr
r
r
r
r
rr rr rr
r
r
rr
rr rr rr rr ff ff
f
GRAINS
r
rr
r
rr
(r)
r
r
r
r
Cat rina
Brazil Brazil Surinam Surinam
Amazonas
Br.
Par na
Guyana Surinam Surinam
Surinam Surinam Guyana
Br.
Surinam Surinam Ven zuela
1335
3898 2943
221
1125
151
M.
L.
3403
&
IANw IANw Stahel L. Pr.
&
F.D.
Notebom Notebom Notebom Notebom
Dietr.
(Spruce)
240
23271
Stahel Heyligers Maguire
H.
&
L.
Spreng.
Sandw.
(Aubl.)
Arn.
ex
Sw.
Sw.
et
A.
Hook,
denta (Aubl.) (Aubl.)
guianesimultifora ultifora raultifora Styrac e glabr tus guianesileprosu
imarubac Quas ia
Styrax
Ternstomia denta puncta
eae
S
cuspidat
Br. Br.
954
4463
Ms.
3535 6742
Surinam Surinam Surinam Pan ma
Br. Br. Br.
Br.
H. H.
de de
349
77
&
&
Guiana
Fr.
Fr.
Surinam unk own Surinam Surinam Brazil Surinam
H. H.
1225
H. H.
L. L.
Guiana
1058 1060
2 680 de de
&
Lindeman Flor. Bret ler Krukof
Amazonas
1016
Stahel BBS
L.
H. H.
684
S
L.
Reitz
USw
Walp.
DC.
(Aubl.)
Br.
Par na Par na
Sta.
&
&
L. L.
7169
M
31
Krukof Stahel
356 9 1076 1015
5636 1195
316
M
317
L.
BAFOG Maguire BAFOG
&
Yw
BBS BBS
L.
Sw.
D.
Jus .) Jus .) Jus .)
500
24 24 de
Schulz Maguire
Par na Par na
2475
H.
7142
Amazonas
Schot
(A. (A. (A.
fanshwei
Theac e
&
Desf.
lancifola
Theoprstace Clavija
ingens
(Spreng.) (Spreng.) (Spreng.)
Spruce
divarcta Verbnace compresa compresa compresa floridula megapotmicaegpota ic megapotmica
Tilace Luehea Vitex
acuminat
Mart.
Desf. Desf. Mez
lancifola lancifola parviflora
Mold. Mold. Mold.
Mart.
candicans
Turcz.
Turcz. Turcz. Duchas .
var.
Warming Warming Warming Aubl. Aubl.
WarmingAubl. Aubl.
Vochysiae Qualea albifora albifora albifora coerulea coerulea ingens rosea rosea
135
B.
J.
H.
TER
WELLE
(1)
Ephedranthus Himatnhus
(1).
Oxandra
(2),
(1),
,
(10)
xylem
secondary
the
Cymbopetalum
grains species
(1),
of
number
.
without
taxa
the
brackets
Neotrpical(betwen
2.
TABLE
(1)
Bocageopsi
Geisoprmu
Trichanter Anona
,
(1)
(4), (3),
Guateria Uno psi
(1),
Pseudocalym
(1),
Terminal
(1).
(1),
Arabidea Potamgenos
(1),
(18), (1), (1),
(2).
(1),
Fresiodnr Panoxdra
(5),
(4).
(2),
Ilex
Discophra
(1),
(4).
(1),
(1),
(2).
Plenckia
(1), (1). (1), (1),
(1),
Lacmel a
(3),
Lagunclari Haripa Weinma i Tapura Dolicarpus
(1),
(1),
(1),
(2). (3).
(1).
Aspido erma Stenoslen Scheflra AnemopagmPhryganocdia Tabeuia
(1),
Em otum
Pinzona
(1),
Rolina Macoubea
(1),
(2),
(1),
(1), (1),
(1),
(5),
Ler tia
Tetrace
(3).
Cydista
Bonafusi Malouetia Tabernmota
(1),
(2).
(1),
(1),
Gonypetalum
Combretum Dicranostyle Lamnoia Davila
(3),
(1).
(3),
,
(10).
(1),
(1),
(13)
(1),
(1).
(6).
Vismia
(4),
(1).
(1),
(1),
(9).
(3),
Dendrobangi
(1),
(2).
Aphelandr Anaxgorea Ambelani Stemadnl Didymopanx Aristolcha Adenocalym Par gonia Stizophylum Cordia Maytenus Cochlspermua Buchenavi Bachris Bonamia Caldcuvia Dichapetlum Curatel Sloanea Lisanthus Gnetum Goupia Hypericum Cal tola Lacistema
(I), (1),
-
Acanth ea Anoace Apocynae
136
Jacrnda
(1),
Fusaea
(6),
Roentg ia
(2),
Hornschuia Xylopia Parhncoia
Couma
(8),
Poraqueiba
(2),
Plumeria
(2), (8).
in
sil ca
Malmea
Dug etia (1),
investgaed)
Musati (1),
-
-
-
-
-
-
-
-
-
-
-
-
-
(1). (1).
-
-
-
-
-
Aquifolace Aralicea Aristolchae Bignoace Boragincea Celastrce Cochlsperma Combretac Corapsitae Convluace Cunoiace Dichapetl Dilenac Elaeocrpea Gentiace Gnetac e Goupiace Hyperica e Icainea Lacistem a
-
SILICA
GRAINS
(3)
(3),
(3),
(1), (5),
Hiraea
(1),
(1),
Loreya
Hetropeis
(2),
(1).
Nepsera
(8),
Henritla Myriaspora
Brosimu
(1),
HelianthosyliPseudolmia
(5),
(5),
Rap nea
(6).
(3),
Virola
Batocrpus Ficus Pour ma Cybianthus
(1),
(4), (1),
(4). (2), (2),
(2),
(1), (20),
Siparuna Bag s a Cousapo Per bea
Clidemia Miconia
(1),
(1),
(1),
(1),
Dolichpterys Tetraperis Henritea Mouria
(1),
(1),
Castila Helicostyli Sor cea Stylogyne
(1),
(1),
(3),
Roucherla Byrsonima Spachea
(11),
Rinorea
(2),
(1), (3),
(1),
(2),
(3),
(4),
(1).
Maquira
(3),
(2),
(1),
(2).
Trymatoc us
Cecropia Weig ltia Campo nesia
Myrcia
Mac irea Tibouchina
(1),
(2),
(1).
(1), (4),
(4),
Osteophl um Conmorpha
(4),
(2),
(1),
(1).
(2).
(5), (1),
CalyptrnhesMltranhes
(1),
(1),
Calycoretus
Marlie a
(1).
Rhizophra
(1),
(1),
Gomidesa
(2),
Calycopus Myrciar
(2).
(1),
Pisonia
(7),
(1),
(1).
(1). (3).
(2), (1),
(1),
Sterculia
(1),
(1),
Gloesprmu
Cyphomandr Herani
(1),
(2),
Casipourea Cestrum Guazma
Securida
(1),
Leonia
(1),
(1),
(5).
Eugenia (1), Neea Ouratea
(2),
(1),
(3). (2),
(3), Solanum
Syz gium
Plina
(2),
Sterigmap lu
(1).
Payp rola
Theobrma
(1),
(1),
(1),
(1).
Trema
(3).
(1).
(2).
Anchieta
(1), (2),
Hebptalum Bunchosia Loph teris Belucia Merian Toc a Molineda Ano carpus Clarisla Naucleopsi Iryanthera Ardisa Blepharoclyx CatingaMyrcianthes Guapira Elvasia Agonadra Pasiflora Piper Moutabe Gouani Anisophylea Brunfelsia Basiloxy n Symplocs Trigonia Ampelocra Amphir ox
-
Linace
-
.
(3)
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
Malpighcea Melastomce Monimace Morace Myristcae Myrsinace Myrtace Nyctagin ea Ochnae Opilace Pasiflorce Piperac e Polygace Rhamncea Rhizoprace Solance Sterculia e Symplocae Trigonace Ulmace Violace
137
B.
J.
H.
TER
Normand
of
WELLE
(1966, 1967) reported
the
frequent
of silica
occurrence
grains
in the ray cells
Q. tricolor.
No silica
grains were
observed in Callisthene
(1), Erisma (5), Salvertia (1),
and
Vochysia
(13).
DISCUSSION
According
to
Amos
the
Chrysobalanaceae)
constant
(94%),
in the
axial
times
in
The
tyloses
the axial
in
remaining
occurrence
is of
no
or
axial
the rays
grains is
a
constant.
The
as
60%
one
not
‘not
a
this
et
fibres is
these results it is
on
the
of the
frequent
is
presence
grains
portion
some-
is
percentages
of silica
are
over
grains
restricted to
of the siliceous
when
grains are
not
species
the
Similarly
where the
concluded that the
restricted
same
grains
are
of
restricted
occurrence
species level, especially
sample
occurrence
of silica
when the
grains
only
a
occurrence
of silica grains in
in many instances the
of silica
presence
it is
of the
not
grains
few, mostly
of such
at
small
the
or
great importance
family
grains
a
genus
may be
at the genus level.
level is very small. Besides the
monotypic families,
show
a constant
grains.
grains
than the
by
means
Grains
may be smooth
ones
of
not
been
with
a
this
type
show
a
or
granular.
Grains with
granular surface. Although
electron
scanning
with
/um). Granular surfaces
character has
the
Silica grains
in other tissues of the
variable.
of all the material
investigated, the
diagnostic feature,
studied
occur
in those
constant
Therefore, though
granular’.
10
(g
and
of the
small
a
grains
of silica grains in those tissues is
occurrence
but also
4%.
in which the
only
of silica
is variable.
tissue.
surface of the
less
was
to
diagnostic importance
occurrence
The
only,
parenchyma
is most
rays
Chrysobalanaceae,
are
the
sum
species (including
follows; ray parenchyma
as
species,
a
promising diagnostic
presence
of the vessels
tyloses
constitute
valuable diagnostic character
In about
valuable
and
very
occurrence
diagnostic importance. However,
only. According
restricted to
are
is
in axial
in the
the fibres
species
the
the various tissues is
over
and fibres
parenchyma
grains
the
a
117 siliceous
species
diagnostic value. Species
to
in these
as
In 110
species
seven
grains is
samples of
tissue and therefore the
one
absolute
parenchyma
their presence
to
more
of tyloses being variable in
it is also of
very constant
silica
grains
than
more
species. However,
to the
or
parenchyma 20%, fibres 4%,
occur
100%.
two
investigated.
were
of silica
occurrence
study,
The distribution of the
85%,
the
(1952)
character. In the present
microscopy,
of surface
are
studied. As
smooth surface
it should be
always
considerable variation
sufficiently
a
this smooth surface
small
or
interpreted
(Plate 2/6-11).
Scurfield
et
as
medium sized
At
present
al. (1974a) and Hirata
al. (1972) also described various types of granular surfaces, it might be of interest to
study
this
phenomenon
more
extensively, especially
with
regard
to
its
possible diagnos-
tic value.
The size of the grains is very variable
138
as
pointed
out
by
ter Welle (1976). The best way
SILICA
Plate
1.—Distribution
cells; 2,
the
ray
Licania
cells;
Mezilaurus
to
make
patterns
majuscula.
4,
Trattinickia
itauba. Globular
a
good
use
of silica
Globular
grains
to oval
demerarae.
in
the rays;
grains
Almost
in
1,
ray
Licania
in
leplostachya.
Globular
restricted
short horizontal
probably
to
bands
found in
the
of
ray
in all
grains
cells; 3, Lecythis davisii.Oblong grains
globular grains,
grains grouped together
of this character is
all
GRAINS
marginal
in
ray
ray
part
of
cells; 5,
cells.
introducing the following
categories:
small
(1)
grains
(2)
grains medium-sized (6-15 /um),
(g
(3)
grains large (g
5
15
/um).
Logically, this classification will apply
to
globular grains only.
139\1
B.
J.
Plate
the
H.
WELLE
2. —Various
ray
lumen
dense
TER
cells;
is
7,
shapes
filled; 8,
inner
Erythroxylon
granular
citrifolium.
quite different
surface is
irregular
from that of
and varies
Silica
grain (arrow); 9,
than that of the
surface is
grains (SEM); 6, Senefeldera macrophylla. Oblong grains
pomifera. Strongly granular
side of the broken
surface is less
140\2
and surfaces of the silica
Achrouteria
from
7;
grains
in
surface
grains
with
Same
sample
6
7;
and
11, Erythroxylon
10,
as
of silica
grain
various
shapes
8, showing
Couratari
nitidum. More
granular to nearly smooth.
an
stellata.
or
less
in
a
in
oval
ray
the
cell. The
cells.
ray
grain
in
a
Oblong grain.
oblong grain
in
ray
The
a
in
entire cell
ray
Note
the
cell,
the
granular
cell. The
SILICA
GRAINS
ACKNOWLEDGEMENTS
I
grateful
am
Gorts-van
useful advice
the
Dr. A.
to
Rijn,
M.
W.
who
Mennega
supervised
this
due
are
for
the botanical names, Mr. L. Y. Th. Westra for correction of
concerning
Thanks
R. A.
acknowledged
version and Dr. W. Berendsen for his assistance with the
English
microscope.
A.
Mrs.
study.
Mrs. M. Jansen-Jacobs and Dr. J. C. Lindeman are
Mr.
to
A.
and Mr. T.
Kuiper
electron
scanning
for
Schipper
the
preparing
photographs.
REFERENCES
Amos,
G. L.
1951.
Some siliceous
Amos,
G. L.
1952.
Silica
P.
Araujo,
A.
P.
Araujo,
A.
&
de
Menon,
P. K.
Leiden
Besson,
bot.
A.
Brazier,
J.
Burgess,
P.
M.
Prison,
1942.
J.
madeiras
133-137.
brasilieras
Estrutura
of Silica
wood
das
madeiras
in
Malayan
woods.
in several woods
of
anatomy
de
39:
de
angiospermas
71-85.
Malay.
Forester 28:284-288.
ofN.S.W. Trop.
Bonnetiaceae,
angiospermas
39: 61-71.
brasilieras
(Pohl) Pittier). Rodriguesia
deposition
cendres et teneur
en
in
1948.
1961.
silice des bois
Malay.
Woods
and
Theaceae,
113:48-54,
Guttiferae.
Bot.
revision
Forester
Ztg.
of the
studies
tropicaux. Agron. trop., Nogent
of hardwoods.
28:
in
15;
Prod. Res. Bull.
Aust. J. scient.
281-292,
1: 44-56.
46.
Res.,
Connaraceae
Ser.
1:
B,
279-302.
297-308.
Contr. U.S. natn.
Humiriaceae.
the
Forest
223-229.
of the Proteaceae.
anatomy
Fragmente.
Anatomical
en
Identification
Sabah timbers.
The wood
A taxonomic
1972.
1889.
Prantl,
De la
W.
Proteaceae.
de
presence
dans le bois du Parinari
Gonggrijp,
12:
II.
Wood
Herb.
anatomy.
35:
J.
25-214.
Elisha
Mitchell
120-136.
A. & K,
E.
das
Estrutura
1974b.
rubescens
1960. Silica
Franklin,
Silica
1961.
E.
scient. Soc. 88:
Engler,
Forester
(this issue).
M.
J.
W.
Caribb.
267.
1974a.
Filho,
(Panopsis
Comparative
1857. Westindische
Cuatrecasas,
Dickison,
1976.
Richesse
F. 1965.
H.
Mattos
de
Lanyon,
D. & G. L.
Chattaway,
CrOger,
Ser.
1946.
Filho.
1965. Guide to distribution
T.
Baretta-Kuipers,
of British Guiana.
(Panopsis sessilifolia (Rid.) Sandw.). Rodriguesia
VIII Proteaceae
R. K, & J. W.
Bamber,
Mattos
de
A.
timbers
timbers. C.S.I.R.O. Bull.
VII Proteaceae
dicotiledoneas
Balan
& A.
de
dicotiledoneas
in
1923.
siliceux
Oliv. et du Dialium
glabra
Over
In: Natiirl.
corpuscules
paalworm
PflFam.
dans les bois
klainei
tegen paalworm
en
tropicaux
Pierre. Bull,
bestand
en
g6n6ral
et
agric. Congo beige
hout.
particulier
en
28;
Indische Mercuur
91-105.
46:
351-353,
369-371.
J.
Gonggrijp,
welke
tegen
Gottwald,
Jb.
87;
Johnson,
family.
H.
1932.
Gegevens
paalworm
& N.
betreffende
bestand
zijn.
Parameswaran,
een
Meded.
1967.
onderzoek
Nederlandsch-Indische
naar
Boschbouwproefstat.
Beitrage
zur
houtsoorten,
25.
Anatomie und
Systematik
der
Quiinaceae.
Bot.
361-381.
L. A. S. & B. G.
Briggs,
Bot. J. Linn. Soc. 70;
Hirata, T.,
cells
W.
den
H.
Saiki,
of certain
Koek-Noorman,
Rubiaceae.
Harada,
woods
1969a.
A
J.
1969b. A
J.
1970.
(Rubiaceae).
Koek-Noorman,
Acta bot. neerl.
J.
21:
1972.
A
1972.
by scanning
contribution
18;
evolution
and classification of
a southern
electron
to the
of
crystals
microscope.
and silica
Bull.
Kyoto
inclusions
Univ.
in
Forests
parenchyma
44:
194-205.
wood
anatomy
of South American
(chiefly Suriname)
to the
wood
anatomy
of
(chiefly Suriname)
South American
18: 377-395.
contribution
Acta bot. neerl.
The
Observations
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