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BLUMEA
46
125-140
(2001)
An annotated key
in
to the genera of
Henk
Missouri
Lauraceae
the flora Malesiana region
Botanical
Garden,
van
der
P.O. Box
Werff
299,
St.
MO
Louis,
USA
63166,
Summary
important component of wet tropical
Lauraceae
are an
Malesiana
region.
poorly
defined and
together
on a
problems
fication
Key
in
are
words.
Their identification
has been
existing keys require
Here
specimen.
generic
a
key
delimitation
are
both
based
forests and
flowers and
almost
discussed and
fruits,
entirely
well
are
in
represented
two factors: several of the
hampered by
on
vegetative
which are,
are
however, rarely present
flowering specimens
characters
the Flora
genera
in
helpful
is
presented,
generic
identi-
listed.
Lauraceae,
genera,
characters.
key, Malesia, flowers, vegetative
Introduction
Lauraceae
are
often
Flora Malesiana
wood
or
are
Lauraceae
to
important component of wet, tropical
an
form
region
exception. Many species
no
commercially exploited. Still,
of
variety
a
remains very difficult.
species
genus or
for
forests and forests in the
locally
are
used for their
identification of
reasons,
Existing keys frequently require
both flowers and fruits and because fruits of Lauraceae need several months before
reaching maturity,
both flowers and fruits
are
Therefore, keys requiring flowers and fruits
rarely present
are
essentially
on
herbarium specimens.
worthless for identification
purposes.
Recently,
the FM
area
some
keys
to
genera found in the FM
were
genera
written. 1
and
area
boiling
up
few flowers when
I also list several fruit and
generic
have
identification.
not
surveyed
the
range of variation in
in the list of fruit and
The brief
the FM
area.
published
for small
suggest; for example
key
based
for which
areas
on
of
flowering
they
material
the number of
characters which
seen
many
Cryptocarya
stamens
seem
species
in
and anther locules.
helpful
indicators for
the smaller genera, I
R.Br, and Litsea Lam. and the full
probably
not
expressed
characters.
for each
genus
larger
species
a
characters in these genera is
vegetative
descriptions given
work outside the
present
parts
include all
necessary tool for identification. I recommend
I have
genera
vegetative
the
a
vegetative
large
Especially
is
counting
Although
not
to
not
Because of the small size of the flowers of Laura-
area.
good dissecting microscope
a
might
it would be useful
thought
for all genera found in the FM
ceae, a
of Lauraceae have been
(Kochummen, 1989; Middleton, 2000). These keys do
apply only
genera may be
of Persea Mill, and
more
to
the
species occurring
variable than the
Caryodaphnopsis
in
descriptions
Airy
Shaw may
126
BLUMEA
have 2-locular
stamens
No.
46,
6 instead of 9 fertile
or
2001
1,
and
stamens
few Chinese
a
species
in Litsea have clustered leaves.
placed
Sterile, fruiting
below. In
pistillate specimens
or
vegetative
some cases
while in other
specimens,
the
—Vol.
characters
generic
has
cases one
to
help
can
the
recognize
key presented
in identification of such
specimen
species,
to
because
visible.
not
are
be identifiedwith the
cannot
fruit characters
or
CHARACTERS
Flowers
For
better
a
of the
understanding
key
and discussions I will
All
the other
a
small
in
with unisexual flowers
species
sex
are
stigma,
without
stipitiform pistillode
flowers the staminodia may have
pistillate
broadened
a
absent. The
of reduction of
degree
and is worth
not
all, genera flowers
are
floral parts:
whorls, which
Tepals:
than the
in
arranged
sometimes
although
they
two
whorl; the reverse, the
outer
are
again opposite
whorl
the
their base 2
at
may form
a
cushion
Endiandra R.Br.)
Pleurothyrium
their base
whorl
III
glands
at
form
enclosing
the
all
due
all
enclosing
to
the
few
stamens
(if
more
and
glands
that
are
either
In
1 locule. The anther locules
most
genera the
(introrse)
stamens
extrorse; in others all
or
are
are
said
exserted. Because in
twisted, it is difficult
to
of the
on
or
and
of
species
neotropical
three) having glands
or
all
glands
stamens
see, not
at
of
have
surprising
small lauraceous flower.
2 locules;
flaps
rarely
the side of the anther
so
char-
do anthers
facing
the
towards the outside of the flower (extrorse).
of whorl I and II have introrse locules, while whorl III is
stamens
genera with umbellate inflorescences,
the anthers
are
facing
4
of whorl
enlarged
some
in small chambers, which open with
have
of the flower
in
(as
stamens
Stamens of
stamens
become
most
a
and
rule
whorl III
tepals,
greatly enlarged
fitted in
a
(I, II, III, IV,
tepals.
small in size and difficultto
are
typically
only
than
of the
as
tepals. Thus,
species
most
In Cinnadenia Kosterm.
glands
pollen
glands
of whorl III
(as in
stamens
the
genera the
the
whorls of
than the inner whorl, is
the inner
most
acteristic for Lauraceae. The anthers have
center
a
may be
shape
inner whorl is
the inner
opposite
cases
in
in four whorls
opposite
In
misinterpretation
stamens.
stamens
produce
a
a
stamens
Nees). Reports of all
mostly
in
the
larger
opposite
whorl of six.
globose glands;
their base, but these
stamens
of
shape
they
following
usually equal
case
whorl
whorl II
and whorl IV
one
surrounding
or
the number of
given
The
are
to
are
arranged
and
inside)
tepals,
outer
tepals
outer
I and II appear
III bear
the
opposite
outer
Stamens:
counted from the outside towards the
of whorl I
or
well documented
not
the
encounters
In that
unequal.
are
(Triadodaphne Kosterm.).
rare
anther
trimerous, with the whorls of floral parts in
center one
very
an
and staminodes is
pistillode
may be absent;
investigating.
In most, but
larger
of
of
with
pistillode
a
pistillode
unisexual.
remnants
of an anther in the
trace
a
find
may
or a
a remnant
threes. From the outside towards the
size,
one
stigma
a
may be threadlike without
tip, they
or
dioecious. In unisexual flowers
are
In staminate flowers
usually present.
describe the
briefly
flower structure of Lauraceae. Flowers of Lauraceae are either bisexual
to
have introrse locules. The latter is the
in which the
stamens
have
these genera the filaments
ascertain if the locules
are
all introrse
or
long
are
not
case
in
filaments and
long
and often
and I have
not
H.
der
van
Werff: An
used this character in the
are
the
lateral. In
case, stamens
ments,
all
the anthers
whorls of
stamens
6
3 fertile
or
same
are
is
stamens.
to
some
(Endiandra, Eusideroxylon
3
only
IV all fertile)
not
are
shorter and
and
stamens
one or
disappear
to
genera it is staminodial, in others it has
stamens
disappear
fila-
long
have
usually
stamens
staminodia. The first whorl
is
com-
and then flowers
have
present, these represent whorl III
are
& Binn. and
Teijsm.
the locules
instance)
number of locules; if this is
space for 12 fertile
enough
reduced
If
127
have fewer locules than those of whorl I and II.
to
not
Lauraceae
for
(in Endiandra,
have the
Sometimes additional whorls
pletely disappeared.
of
genera
exserted from the flower. When the
whorl IV, the innermost whorl; in
only
the
to
(whorls I, II, III,
stamens
are
included in the flower, there
more
key
some cases
stamens
of whorl III tend
Flowers with 12 fertile
so
In
key.
species
most
annotated
An
Triadodaphne).
of this
example
trend towards reduction of number of stamens in smaller flowers is found in the genera
Kosterm.
Potoxylon
and
Eusideroxylon.
all
By
these
accounts
genera
closely
are
related; Potoxylon has larger flowers and 9 fertile stamens, while Eusideroxylon
much smaller flowers with 3 fertile
and with
one
unprotected
a
occupies
an
plate-like
completely
Inflorescence
its base,
or
is seated in
Eusideroxylon
and Poto-
receptacle. Cryptocarya
in flowers of this genus the ovary
is enclosed in
the fruits
technically superior;
receptacular
tube and
thus inferior.
are
types
The inflorescences of the Lauraceae
(excluding
three main types (Van der Werff& Richter,
to one
In
at
and thus hemi-inferior; fruits
receptacle
fused with it and thus
not
and the fruit is seated
superior
cup-shaped.
or
enclosed in and fused with the
are
is
enclosed in and fused with the
position;
tube, but
ovary
has
unilocular
unicarpellate,
persistent tepals
embedded in the
partially
are
receptacular
however
be shallow,
intermediate
the
exceptions
sometimes with
pedicel,
can
the ovary is
in these genera
the
the
which
cupule
xylon
ovule. With few
on
The ovary is
stamens.
of these groups, because in
reduced while
a
few
species
or
some cases
genera
Cassytha L.)
be divided into
can
be
assigned
inflorescences have become
strongly
1996).
Not all inflorescences
have inflorescences which
can
simply
do
not
match
of the three types.
any
In the first type the flowers
are
arranged
in
when young,
umbels, which,
in decussate bracts. All flowers with this type of inflorescence
bels
can
be
single
in the axils of leaves
shortshoots. Umbels
can
be sessile
or
or
bracts
or
grow into
can
are
be
in
In
bracts
a
the shortshoot has
normal
easily
pairs,
vegetative
a raceme
whereas the
tepals
are
not
of
single
Inflorescences of the second type
by
are
a
dichasium
racemose
are
are
reduced
to
a
um-
leafless
arranged along
a
contrast
flowering,
can
the decussate bracts,
single flower;
case
the umbels
the presence of
the inflorescence consists of
flowers.
are
paniculate-cymose.
These inflorescences
in dichasia. The
always strictly opposite (Fig. 1).
through
are
surrounding
branched, with the flowers ultimately arranged
flowers of such
rescences
arranged along
of umbels, but in
still covered
enclosed
in whorls of three.
Blume the umbels
1-flowered umbels and
are
unisexual. The
terminal bud, which, after
leafy twig. Young umbels,
the base of the flower indicates that in this
repeatedly
be
mistaken for flower buds. However, the bracts
Iteadaphne
at
a
can
stalked. When the umbels
leafless shortshoot, the whole structure looks like
to a raceme,
they
are
reduction of the lateral
axes.
are
lateral
Sometimes the inflo-
BLUMEA
128
Vol.
46,
Fig.
No.
2001
1,
1. Detail of
cencewith
a
paniculate-racemose
the lateral
flowers of the
inflores-
cymes
strictly
opposite (type 2).
Inflorescences of the third type
the ultimate cymes
alternate
short
always
not
also
not
paniculate. However,
especially
to see,
easy
not
of Beilschmiedia Nees and Endiandra tend
makes it easier
racemose
to
recognize
this inflorescence
inflorescence with
rarely
three
or
Two genera (
area
Actinodaphne
Nees and
in
not
to
have
has
together
by
in
somewhat
with very
Cryptocarya).
a
on
predominantly
the basal lateral
reduction of the lateral
type.
Cinnadenia)
occur
in the Flora
Malesiana
which have inflorescences unlike the three types described above. The type
ofActinodaphne,
as
well
as
several other
which
longer pedicels
type. Potoxylon
four flowers
are
species
uncommon
branches of the inflorescence. This inflorescence is derived
branches from the inflorescence of the third
the lateral flowers of
and instead
strictly opposite,
and subsessile flowers (a condition
pedicels
Species
This is
(Fig. 2).
are
frequently
are
species placed
in that genus, has
a
species
paniculate
inflorescence which is enclosed in bracts when very young. These bracts do not appear
to
be decussate, but
inflorescence,
rescences
1-2
cm
in A.
long
are
but leave
alternate.
a
pilosa (Lour.)
and
finally
pseudo-umbellate
They
cluster of
there
Merr.
are
fall off very
scars
are
several
to
at
10
cm
long,
species placed
inflorescence, this still with bract
ference between the
early
Actinodaphne species
in the
development
of the
the very base of the inflorescence. Inflo-
with
a
in A. malabarica Balakr.
in
Actinodaphne
scars
at
the
very
pseudo-umbellate
with
a
only
sessile
base. The dif-
inflorescence and
H.
van
der
Werff: An
annotated
key
to
Fig.
the
genera
2. Detail
of
a
of
Lauraceae
paniculate
129
inflorescence
the ultimate flowers not arranged in
strict
with
cymes
(type 3).
Litsea
species
with sessile umbels is difficult
clustered leaves.
area
Fortunately,
have alternate
or
the great
to see,
majority (if
subopposite leaves,
while
are
with
species
of Litsea in the FM
all) species
Actinodaphne species generally
whorled leaves. The inflorescences of Cinnadenia are
niculate, the flowers (unisexual)
in Litsea
especially
not
paniculate
racemosely arranged along
or racemose.
the
secondary
have
If
pa-
axes
of
the inflorescences.
Fruit and
fruit and
mind
some
floral characters
vegetative
though
that there
characters
vegetative
Although
of prime
may
be many
I have used the
usually
term
exceptions
vegetative
perulate
some
internodes.
importance
for the determinationof genera,
times also be very
only give
to
an
helpful.
indication of
One should
keep
generic assignment
these indications. Still,
it is worthwhile
in
and
to
list
characters.
buds for
fall off after the buds
twigs. Only rarely (in
the base of the
at
can
that these characters
of these fruit and
bracts
are
characters
vegetative
elongate
Neolitsea
buds covered
by
bracts. These
and leave distinct clusters of scars
species)
do the bracts
persist
as a
on
the
short sheath
at
BLUMEA
130
Vol.
In triveined leaves the main lateral veins
the lamina, while in
No.
46,
depart
2001
1,
from the midrib at the very base of
leaves these veins leave the midrib
tripliveined
a
little above the
base of the lamina.
-
Fruits enclosed in the floral tube and with the
fruit:
large,
Fruits
less
generally
seated in
a
than 5
cupshaped
of the fallen tepals
scars
Potoxylon. Eusideroxylon
rather slender fruits (more than 5
has fruits
-
and
Cryptocarya, Eusideroxylon
cm
long,
2-3
cm
on
top of the
Potoxylon
have
wide) whereas Cryptocarya
long.
cm
Neolitsea Merr. and
bowlshaped cupule: Litsea,
or
and
Actinodaphne.
-
-
-
Fruits with
persistent, spreading
Fruits with
persistent, appressed tepals
Fruits
seated
unprotected
to
the
on
reflexed
tepals
at
the base: Persea.
the base: Phoebe Nees.
at
pedicel,
without
persistent tepals
or
a
cupule:
Alseodaphne Nees, Beilschmiedia, Brassiodendron C.K. Allen, Dehaasia Blume,
Endiandra.
-
with
Twigs
pale,
whitish bark, this
with the dark
contrasting
petioles: Alseodaphne,
Dehaasia.
-
Leaves clustered, without
Alseodaphne, Dehaasia,
-
Leaves clustered, with
Leaves
clusters
or
of
the branches:
along
scars
buds and clusters of
perulate
scars
along
thebranches: Actino-
Persea.
daphne, Neolitsea,
-
buds
perulate
Phoebe.
opposite: Caryodaphnopsis,
CinnamomumSchaeff. p.p.,
Iteadaphne Blume,
Litsea p.p.
-
-
-
-
-
Leaves triveined
tripliveined
or
opposite:
Cinnamomum.
Caryodaphnopsis,
triveined, alternate: Lindera Thunb. p.p., Neocinnamomum H. Liu.
Leaves
tripliveined,
Terminalbuds
alternate:
working
comment
p.p.,
Dehaasia
A
with Lauraceae,
briefly
scarcity
on
of good,
Lauraceae
collections
good
ments
(for
IN
two
flowering
are
problems
are
trees
soon
Caryodaph-
become apparent and I
and have small,
often among the
ten
(Hyland, 1989)
1986, in which
these collections
in which
It took
nearly
more
a team
together.
to
want
to
the fact that many
flowers. Even
inconspicuous
most common
few. This is reflected in
great majority of species.
bring
p.p.,
STUDYING LAURACEAE
from fewer than 10 collections). A welcome
to
Alseodaphne
collections. No doubt this is due
instance Kostermans,
in Australia
p.p.,
Neolitsea
p.p., Litsea p.p.,
p.p.
those:
Lauraceae grow into rather tall
though
Lindera p.p., Litsea p.p., Neolitsea, Persea.
glaucous: Actinodaphne
PROBLEMS
When
p.p., Lindera p.p.
Cryptocarya
perulate: Actinodaphne,
Lower leaf surface
nopsis, Cryptocarya
1.
and
Leaves
families in forest
all revisions
most
exception
species
or
floristic
treated
are
plots,
treat-
known
is the Revision of Lauraceae
than 20 collections
were
studied of the
of dedicated field collectors several years
For the FM
area,
good flowering
collections
are
very much needed.
2.
The poor delimitation of genera. In
Werff& Richter,
tem:
earlier classifications
(reviewed
1996) generic delimitationwas largely based
on a
in Van der
numerical sys-
number of fertile stamens, number of anther locules, combined with inflores-
H.
these characters results in
with its
mostly
6 2-locular
in
use
forming
delimitation will
groups of
it
I would
mately
the
relationships
not
yet
at
acters.
One
I
was
premature
taking
into
unable to
recognize
species
not
key.
In the other
case,
our
understanding
More and better collections
THE GENERA
la. Herbaceous
b. Erect
a
a
a
parasitic
woody plants
sometimes
two
1989)
allows
or
Alseodaphne
needed
to
was
pointed
solve these
IN THE
vine with leaves reduced
regular
based
Nothaphoebe
OF LAURACEAE
with
Nothaphoebe,
to
floral char-
of the two, I include
and have therefore included
two
of this group, as
are
on
character used in earlier
separation
and
Alseodaphne
genera
in the number of anther cells,
along
on
out
did
not
s.l. and, with
of new
collections has
Kochummen
(1989).
problems.
FLORA MALESIANA AREA
to
. .
minute bracts
6.
Cassytha
2
or
rarely (in
some
or
bracts,
Actinodaphne species
3
paniculate, rarely
8
racemose
dimerous
4
b. Flowers trimerous
5
Inflorescences umbellate; terminal buds
b. Inflorescences
paniculate;
terminal buds
perulate;
not
stamens
surrounded
6
18. Neolitsea
....
by bracts;
stamens
18-
7. Cinnadenia
32
5a. Leaves whorled
b. Leaves alternate,
hnd
Nothaphoebe
description
fruiting
by
I
leaves
leafless shortshoots
b. Flowers bisexual, inflorescences
4a.
Ulti-
classification reflect-
a
Cinnadenia) paniculate
3a. Flowers
so
on
towards the recog-
2a. Flowers unisexual, arranged in umbels, these single in the axils of leaves
and
still
monophyletic.
are
genera and
groups
are
classification based
practical key
if the genera
or
likely
way of
practical
of Lauraceae
species
Dehaasia is very similar
Dehaasia
Alseodaphne,
KEY TO
make
want
develop
to
will
monophyletic
and
poly-
forming monophyletic
and Phoebe; because the fruit
difference between the
helped
Species
stamens
genera
of
recognition
generally accepted
two
instance Kochummen,
Alseodaphne. Vegetatively,
in
to
try
account
differences in flower or fruit between the
only
to
rec-
between these genera, but the information needed for this is
is Persea
case
them both in my
the
Because
taxa.
be
Van der Werff & Richter,
the
are
species
hand.
publications (for
in
of
can
Caryodaphnopsis,
monophyletic
than those who
prefer to recognize monophyletic
ing
cases
between
seems
nition of genera without
result in
that genera
require
and I have therefore tried
relationships
is
example
within
One of the main aims of
species.
relationships
express
ordening
and distinct wood anatomy.
examples (see
likely
generic circumscriptions
incompletely known,
two
This and other
genera. Those who
have different
131
have 9 4-locular stamens, but sometimes 9 2-locular
stamens.
generic
paraphyletic
In
genera and allows the
recognizable
indicate that androecial characters vary
1996)
Lauraceae
of
genera
bisexual), and fruit type. A classification based
or
opposite leaves, strongly unequal tepals
or even
to
to the
key
other characters than those listed above. An
of that genus
their
annotated
such, it is practical. Some genera, however,
groups. As
recognizable
ognized by
is
der Werff: An
type, flowers (unisexual
cence
on
in
van
1.
rarely subopposite
or
opposite
Actinodaphne
6
132
BLUMEA
Vol.
No.
46,
1, 2001
6a. Anthers 4-locular
16. Litsea
7
b. Anthers 2-locular
7a. Umbels 1 -flowered; leaves
subopposite
or
14.
alternate
b. Umbels with several flowers; leaves alternate
Iteadaphne
15. Lindera
8a. Anthers 2-locular
9
b. Anthers 4-locular
17
9a. Fertile
stamens
3
b. Fertile
stamens
6
10a.
Tepals equal
10
9
or
the
or
11
three
outer
smaller than the inner three
slightly
ll.Endiandra
b.
Tepals
the
unequal,
flowers
apparently
1 la. Fertile
stamens
6
b. Fertile
stamens
9
12a.
Tepals longer
outer
with
three much
only
three
larger
and
covering
the inner three, thus
22.
tepals
12
13
than stamens,
included in the flower; anther cells lateral,
stamens
4. Brassiodendron
slit-like
b.
shorter than
Tepals
Triadodaphne
stamens
stamens,
exserted;
anther cells
extrorse,
13.
roundish
13a. Leaves
opposite
b. Leaves alternate
14a. Outer
or
subopposite
or
clustered
tepals clearly
smaller
small,
Hexapora
8. Cinnamomum
14
half the
(about
size)
than the inner
ones
10. Dehaasia
b.
Tepals equal
or
15a. Flowers with
a
nearly
15
so
tubular
deep,
receptacle,
this
enclosing
the ovary and later the
fruit
9.
b. Flowers with
quently
a
shallow
with small,
persistent tepals
16a. Lateral flowers of the ultimate
light-colored
and
the fruit
receptacle,
contrasting
at
cymes
same
stamens
3
b. Fertile
stamens
9
opposite,
b. Leaves alternate,
19a.
b.
the
with the much darker
usually
...
twigs
and
petioles
12.
Eusideroxylon
18
tri-
or
19
tripliveined
pinnately
or
triveined
20
8. Cinnamomum
Tepals unequal,
the
pinnately
21a. Inflorescences
b. Inflorescences
outer
tepals
much less than half the inner
ones
Caryodaphnopsis
17. Neocinnamomum
veined
racemose
21
or
nearly
21.
so
the
outer ones
about half
as
long
as
the inner
ones
2.
or
nearly
so
Potoxylon
22
paniculate
Tepals unequal,
Tepals equal
twigs usually
3. Beilschmiedia
20a. Leaves triveined
b.
infre-
10. Dehaasia
petioles
subopposite;
5.
22a.
or
16
Tepals equal
b. Leaves
Cryptocarya
pedicel
bark of the
color
17a. Fertile
18a. Leaves
on
the base
strictly opposite;
b. Lateral flowers of the ultimate cymes
about the
unprotected
Alseodaphne
23
H.
23a.
van
der
deciduous
Tepals
Werff: An
annotated
persisting
or
as
key
the
to
small scales
at
Lauraceae
of
genera
133
the base of the fruit
2.
b.
24a.
b.
1.
Tepals persisting
Tepals clasping
Inflorescences
19. Persea
Nees
paniculate
umbellate with clusters of bract
or
stamens
scars
at
the
very
base.
9, each with 4 locules, tepals equal. Fruit seated
small, flat cupule. Leaves whorled, usually pinnately veined, sometimes glaucous
below. Terminal buds
immediately
perulate
terminal buds.
whorled leaves, but differ from
having
clusters of
usually perulate;
above the whorls of leaves. Best
whorled leaves, and
in
20. Phoebe
reflexed in fruit
to
Flowers triinerous, unisexual,
on a
24
the base of the fruit
Tepals spreading
Actinodaphne
Alseodaphne
fruiting stage, conspicuous
in
the umbels
generally
the inflorescence. About 100
present
scars
its
recognized by
Some
on
twigs
rather
large,
of Litsea may also have
species
with umbellate inflorescences
Actinodaphne species
stalked and in the absence of bract
Asia with
species, tropical
the older
usually
few
a
scars at
the base of
in southern
species
China.
2.
Alseodaphne
(including Nothaphoebe)
Nees
Inflorescences of type 2. Flowers trimerous, bisexual,
tepals unequal
and
to
Fruit seated
equal.
brightly colored,
or
small
unprotected
tepals
persisting
in several
this
light-colored bark,
About 90
with the darker
Various authors have tried to separate
at
pedicels thick, fleshy
vs.
in fruit
tepals
vs.
equal
vs.
unequal
most
a
club-shaped;
Bentham
and
or
(1880)
qualitative,
and Kostermans
subgenera
opinion
and
a
tepals
Tropical
one was
was
as a
distinct
American
vs.
fleshy
and
only
have
deciduous
tepals
tepals persistent
vs.
quantitative (tepals
subequal
tepals
and
and the
intensively
as
Nothaphoebe
Alseodaphne
in his
Kostermans
Nothaphoebe
more
and
Alseodaphne
key Nothaphoebe
valid genera. Even
than
Nothaphoebe.
has been
separating Alseodaphne
as
the
type
published,
and
as
in Persea
(1973a) changed
anybody else,
I found in BO
annotated in 1960 as the type of Nothaphoebe
names
stami-
vs.
tepals equal; fruiting
ones are
genus (however,
genera).
annotated in 1992
Kosterm. Neither of these
the difficulties of
not
species
treated
clear concept of Alseodaphne and
Kosterm., the other
perulate.
vague).
(1957)
recognized Alseodaphne
of Clemens 29777;
unequal
the other
but several
Kostermans studied Asian Lauraceae
developed
twigs,
Characters
Alseodaphne.
of Persea; Hutchinson (1964) included
recognized Nothaphoebe
included among the
not
conspicuous, cordate-sagittate
in fruit. Of these characters,
distinction becomes therefore rather
sections
plate-like
a
of the
conspicuous,
Terminalbuds
from
Nothaphoebe
qualitative difference;
looks
tips
often with
Twigs
petioles.
fruiting pedicels cylindrical,
persistent
deciduous in fruit is
this sometimes swollen
Asia.
used for this include staminodia of whorl IV
nodia much smaller,
pedicel,
the base of the fruit;
at
below.
species glaucous
contrasting
species, tropical
9, each with 4 locules,
stamens
the
Leaves clustered near the
cupule rarely present (A. insignis Gamble).
pinnately veined,
on
of Alseodaphne
though
he
two
is
his
never
sheets
clemensiae
kinabaluensis
but these annotations underscore
Nothaphoebe.
It
seems
therefore best
BLUMEA
134
to
include
needed
to
in
Nothaphoebe
Vol.
and would
placed
Asia. Good
is needed for their
with
in fruit
of all
study
very similar
had
but
towards
s.l.
tropical
three undescribed species
vegetative
characters
clustered leaves which
clearly point
pointing
Alseodaphne
to
been collected in
they
species (apparently
description,
involved is
species
several genera. It is worth
are
Alseodaphne
pale bark, dark, long petioles, large,
and thickened pedicels
a
one or
Neotropics
material of these
flowering
involved)
are
be described in
certainly
in
2001
1,
Alseodaphne although
determine if all shouldbe
that several collections from the
out
No.
46,
are
(thick twigs
glaucous below)
Alseodaphne.
3. Beilschmiedia Nees
Inflorescence
tepals equal,
seated
I have
paniculate (type 3).
erect
unprotected
not
seen
About 250
This
Flowers trimerous, bisexual,
stamens
9, 2-locular;
anthesis, deciduous in older flowers. Fruit without
at
the
on
pedicel.
with
species
Leaves alternate
glaucous
or
cupule,
a
veined.
subopposite, pinnately
lower leaf surfaces. Terminal buds
perulate.
not
species, pantropical.
confused with
has been
genus
with 9 2-locular
stamens.
The best
Dehaasia, which has also bisexual flowers
difference between the
is the inflorescence
two
type; type 3 in Beilschmiedia, type 2 in Dehaasia. Dehaasia species have also frequent-
ly
clustered leaves and
the
note
with
twigs
under Endiandra. A
persistent (but small) tepals
placed
in
distinct genus,
a
contrasting
species placed
the base of the fruit; such
at
in New Guinea. Whether these
be
bark and
pale
few Australian
species
requires
dark
species
can
be
expected
best included in Beilschmiedia
are
further
See also
petioles.
in Beilschmiedia have
also
should
or
study.
4. Brassiodendron C.K. Allen
Inflorescence
stamens
thesis. Fruit without
a
6, 2-locular, tepals equal, half-erect
seated
cupule,
lower leaf surface
pinnately veined,
all from Australia with
These
reduction sometimes appearing
paniculate (type 3), by
trimerous, bisexual,
do
species
not
one
not
species
form
unprotected
a
Brassiodendron
a
closely
5.
related
as
to
key
which is difficult
Airy
cupule, unprotected
group and
to use
sole
on
the
one
pedicel.
Caryodaphnopsis
species
is
strongly unequal tepals.
having
(type 2).
outer
below. Terminal buds
but differ in
species,
Hyland (1989) placed
them
and for practical
species
reasons
occurring
I
recognize
in New Guinea is
Shaw
paniculate-cymose
tropical America,
6
Endiandra.
4-locular; tepals unequal, the
glaucous
perulate.
solutionmay well express theirrelationships
distinct genus. The
Caryodaphnopsis
Inflorescence
not
at an-
Leaves alternate,
pedicel.
Terminalbuds
glaucous.
in Endiandraand Beilschmiedia. Hyland's
better, but results in
the
Flowers
spreading
also in New Guinea.
homogeneous
a
on
racemose.
or
easily
not
Flowers trimerous, bisexual,
3 about 1/5
Leaves
area
recognized
long
opposite,
perulate.
in the FM
as
by
as
tri-
About 15
flowers with
equal tepals.
or
tripliveined,
species
in
(Philippines, Borneo,
its
opposite
Cinnamomumspecies often have
stamens
9,
the inner 3. Fruit without
more or
tropical
New
a
less
Asia and
Guinea).
leaves and flowers
with
opposite, tripliveined leaves,
H.
6.
Inflorescence
the
der Werff: An
annotated
key
the
to
genera
Flowers bisexual, trimerous,
a raceme.
outer
in Australia,
This herbaceous,
perulate.
not
enlarged
About 20
un-
floral
species,
species pantropical.
one
parasitic
vine
be confused with any other genus of Laura-
cannot
sometimes it is confused with Cuscuta
ceae;
135
9, 2-locular; tepals
stamens
3 much smaller than the inner 3. Fruit enclosed in the
tube. Leaves alternate, minute, scale-like. Terminal buds
mostly
of Lauraceae
L.
Cassytha
equal,
van
(Convolvulaceae).
7. Cinnadenia Kosterm.
Inflorescence
a
plate-like cupule.
rarely
collected
The
with 9
Leaves alternate,
genus
of two
which differs in
(18-32
stamens
type species
having
as
having
in C.
C.
as
a
a
seen an
synonym
flowers.
malayana
stamens
isotype
are
malayana
and found it
seen were
smaller towards the
have 4
center
Kochummen
(1989)
dimerous, but Kostermans
suggesting
genus;
species (and
to
A
Kostermans (1973b) described the
that the flowers
which Kostermans
(Hook.f.) Kosterm.,
known
poorly
a
small,
inflorescence is Actino-
Long (1984).
liyuyingii H. Liu,
of Cinnadenia
paniculata
Cinnadenia is
on a
perulate.
not
terminal buds, and flowers
1 have
not
are
(1973b)
and this has dimaterial of
seen
thus the concept of Cinnadenia in
Kochummen's (1989) observations. Dr. J. Rohwer
on
flower of C.
progressively
did
he had
paniculate
a
in whorls of 3,
of Litsea
(see discussion),
Malaysia.
perulate
malayana Kosterm.).
and base my view of this
malayana
this article)
in Peninsular
as
dimerous
veined. Terminal buds
whorled leaves,
trimerous flowers,
(1977) mentioned that the
trimerous. I have
merous
one
or
4-locular. Fruit seated
(9-32),
pinnately
species,
mentioned that the flowers of C.
treated
variable
stamens
other genus with unisexual flowers and
only
daphne,
Flowers unisexual, trimerous
panicle.
number of
tepals equal,
tepals
and 28
(pers. comm.)
stamens
checked
which became
of the flower.
8. Cinnamomum Schaeff.
Inflorescences
paniculate-cymose (type 2).
4-locular; tepals equal, half-erect
or
without
persistent tepals
tripliveined
About 350
or
Species
nately
Jaya
alternate and
species, mostly
Oceania and
or
tropical
with
in
tepals partly persistent.
pinnately veined; rarely
and
tropical
tripliveined
leaves
perulate
usually
lack
buds. A few
most
pinnately
veined leaves and
tepals equal,
small
but also
stamens
cupule,
Leaves either
perulate buds,
species
9,
this with
opposite
and
tripliveined.
in Australia,
present
species
Philippines
with
pin-
and Irian
1986) instead of 4-locular anthers. The
Cinnamomum species in the FM
perulate
while
from the
leaves combined with flowers with
fication of
Cryptocarya
a
leaves alternate and
subtropical Asia,
have 9 2-locular anthers (Kostermans,
Inflorescences
Flowers trimerous, bisexual,
anthesis. Fruit seated in
America.
veined leaves have
opposite, tripliveined
9.
the
at
equal tepals
area; one or two
allow
species
easy
identi-
have alternate,
terminal buds.
R.Br.
paniculate (type 3).
half-erect to
Flowers trimerous,
spreading
at
anthesis. Fruit
bisexual,
entirely
stamens
9, 2-locular;
enclosed in the
enlarged
136
BLUMEA
floral tube,
at
the
with small
tip
ribs. Leaves alternate,
perulate.
Several
species
have
among Lauraceae, but this
are not rare.
species
in
A
monotypic
in
few
a
somewhat
also
leaf
uncommon
not
shape
Glaucous lower leaf surfaces
with the
species, pantropical,
their 9
genera,
2-locular
Eusideroxylon
and
stamens
and
floral tube
deep
of
majority
larger (more
than 5
distinctive.
are
have also fruits
Potoxylon,
floral tubes, but both have much
enlarged
an
longitudinal
Terminal buds
species tripliveined.
in Litsea.
occur
with
frequently
oblong leaves,
than 300
more
2001
1,
Asia.
tropical
The flowers with
Two
in
or
might
genus of
large
No.
46,
from the fallen tepals,
scars
pinnately
Vol.
fruits.
long)
cm
enclosed
fully
10. Dehaasia Blume
Inflorescences
deciduous
or
but
persistent,
sometimes with very small,
species
not
trimerous, bisexual,
3 much smaller than the inner
outer
in fruit. Fruit
enlarged,
persistent tepals
sometimes
whitish bark, which
species).
Terminal buds
Dehaasia is
closely
related
to
and without
assigning
stamens
a
the
it is
specimen
pedicel,
in
some
a
pale
to
in Alseodaphne
Asia.
in
only
indistinguishable
either of these
to
have
twigs
case
from which it differs
Vegetatively,
the
equal,
of the branches,
tips
is also the
species, tropical
Alseodaphne,
2-locular instead of 4-locular anthers.
on
9,
stamens
almost
to
fruiting pedicel
species
some
petioles (this
About 35
perulate.
not
below. In
glaucous
with the dark
contrasts
3)
unprotected
the base, the
at
swollen and colored. Leaves alternate, clustered near the
pinnately veined,
daphne
Flowers
paniculate-cymose (type 2).
2-locular; tepals unequal (the
having
from Alseois
genera
guess.
a
11. Endiandra R.Br.
Inflorescences
paniculate (type 3).
Flowers trimerous, bisexual,
tepals equal, rarely subequal (with
ones),
half-erect
to
spreading
at
the
tepals slightly
outer
anthesis. Fruit
unprotected
deciduous in fruit. Leaves alternate,
pinnately veined,
buds
the
not
greatly enlarged
About 100
area
can
12.
only
two
be
not
disc
not
the
of the
surrounding
the
to
the
the
pedicel,
glaucous
the base
3, 2-locular;
below. Terminal
stamens
stamens
erect at
with small
the
tepals
outer
either
genus
anthesis. Fruit
Vegetatively
cences
entirely
from fallen
below. Terminal buds
not
tepals.
perulate.
very similar
and flowers with
pistil.
to
with
much
large
larger
stamens:
than the inner
Triado-
ones,
and
staminodia (representing whorl
Beilschmiedia, and fruiting specimens
certainty.
& Binn.
laxly paniculate (type 3).
scars
become
and
Flowers
trimerous, bisexual,
stamens
locular; staminodes representing whorl I and II well developed; tepals equal,
less
inner
tepals
Australia and the Pacific.
Endiandra is similar
Teijsm.
at
on
other genera have flowers with 3 fertile
having
assigned
Eusideroxylon
Inflorescence
a
which has 4-locular stamens and
II). Vegetatively
often
glands
united, forming
are
which differs in
Eusideroxylon,
I and
and
species
some
species, tropical Asia,
In the FM
daphne
In
perulate.
stamens
smaller than
to
only
enclosed in the
accrescent
Leaves alternate,
Wood very hard. 1
Potoxylon,
three fertile
species,
but differs in its
stamens.
floral tube,
pinnately veined,
not
3, 4-
more or
at
the
tip
glaucous
Borneo and Sumatra.
laxly
branched inflores-
H.
13.
der Werff: An
van
annotated
to the
key
of Lauraceae
genera
137
Hexapora Hook.f.
Inflorescence
paniculate (type 3)
6,2-locular; tepals equal,
tepals
and
stamens
at
erect at
the base
clustered, pinnately veined,
typic
genus,
rarely
(Kochummen,
recognized
whorl I and II and
by
Flowers
trimerous, bisexual;
known from
only
its 6 2-locular
by
the
1989).
stamens.
Fruit with
Leaves alternate
below. Terminal buds
glaucous
not
collected and
is best
Hexapora
senting
or racemose.
anthesis, shorter than the
somewhat
or
with round locules repre-
locules, which is very unusual for
extrorse
14.
long,
slit-like anther cells. Neither
tepals
Inflorescence umbellate, the umbels 1-flowered,
shortshoot. Flowers trimerous, unisexual;
Fruit
not
seen,
said
genus
be seated
to
opposite, pinnately veined,
typic
not
glaucous
from Java and Sumatra,
S Chinaand NE India has also been
This
species
has also
on
a
few
related
related to I. caudata and that the buds of L.
I expect that
umbels
I.
to
confusa,
lated
to
a
more
thorough study
opposite
or
A
perulate.
subsub-
mono-
differing
level. A
generic
on
in its
species
stamens
along
leafless short-
leaves. It is
likely
closely
young, bracteate inflorescences.
are
find that
from
[I. caudata (Nees) H.W. Li].
tripliveined
or
the reduction of multi-flowered
1-flowered umbels in Lindera has taken
with its
or
or
Kosterm. (Kostermans, 1992) is
spicata
will
not
Lindera, but
to
in Iteadaphne
spicata
opposite
1981).
recognition
placed
slender, leafless
a
Leaves
cupule.
1-floweredumbels with 2-celled
described Lindera
recently
together along
below. Terminal buds
shoots, but differs in having alternate, strongly trithat the
persist
lateral
are
in fruit.
6-9, 2-locular, tepals equal
small
(Richter,
certain Iteadaphne deserves
am not
a
stamens
closely
1-flowered umbels and wood anatomy
I
nor stamens
stamens
Blume
Iteadaphne
equal.
mono-
Malaysia.
of whorl I and II. Brassiodendron has also 6 2-locular stamens, but these
and have
A
perulate.
Peninsular
Penang,
stamens
not
stamens
persistent
place
subopposite, pinnately
more
than
veined leaves is
I. caudata and L. spicata,, which have alternate, tri-
or
once
and that
closely
not
re-
leaves.
tripliveined
15. Lindera Thunb.
Inflorescence umbellate, umbels with several flowers,
of leaves. Flowers
nearly
so.
trimerous, unisexual;
Fruit with
or
without
a
small
stamens
9
along
or more,
a
shortshoot
Leaves alternate,
cupule.
pinnately
veined, the lower surface sometimes glaucous. Terminal buds perulate
of about 100
in
species, mostly
tropical
and
temperate N America; relatively few species
Lindera differs from Litsea
specimens
or
specimens
with
only
subtropical
in the FM
in axils
or
2-locular, tepals equal
Asia with
a
or not.
few
or
A genus
species
flowers
are
in
area.
in its 2-locular (not 4-locular) anthers and
pistillate
or
tripli-
difficult
to
determine
to
fruiting
genus.
16. Litsea Lam.
Inflorescence umbellate, umbels with several flowers,
leaves. Flowers
lacking.
trimerous,
Fruit with
a
small
unisexual;
to
rather
stamens
along
shortshoots
or
in axils of
usually 9, 4-locular, tepals equal
large cupule.
Leaves alternate
or
or
rarely opposite,
138
BLUMEA
pinnately veined,
more
genus,
sometimes
than 300
Vol.
glaucous
in
species, mostly
No.
46,
1,
2001
below. Terminal buds
perulate
A
not.
or
large
tropical Asia, also present in Oceania, Australia
and in N and C America.
Litsea is
characterized
4-locular anthers. It is
area,
the inflorescences
bearing
shortshoot
genus.
leaves in the FM
tripliveined
shoots
genus in the FM
variation in the
morphological
or
its umbellate inflorescences, trimerous flowers
by
large
a
illogical)
in
some
can
species
of umbels; however, there is
raceme
'raceme' is
twig bearing
a
Although
I would
be
and
area
I have
be
not
not
always
may expect
if
they
10 cm, and
to
and
of
great deal
with clustered
exist. The short-
making
terminal bud
vegetative
a
the
term
shoot looks much like
umbel-bearing
a
species
seen
surprised
quite long (up
and the
one
indicating
a
the
umbels.
17. Neocinnamomum H. Liu
Inflorescence
or
paniculate-cymose,
inflorescence reduced
to a
4-locular, tepals equal, half-erect
species
not
perulate.
anthesis. Fruit
strongly
A genus of 6
tri-
species
or
in
on an
stamens
9,
obconical, swollen pedicel,
not
tripliveined,
tropical Asia,
below.
glaucous
with 1
poorly
known
and the swollen
pedicel
from Sumatra.
The fascicled flowers, triwith
fascicle. Flowers trimerous, bisexual;
at
Leaves alternate,
tepals persistent.
Terminal buds
the cymes often sessile and the flowers in fascicles
single
persistent tepals
or
tripliveined,
in fruit make
alternate
leaves,
rather easy.
recognition
18. Neolitsea Merr.
Inflorescence umbellate, umbels with several flowers,
along
shortshoot. Flowers dimerous, unisexual;
a
4, spreading
at
anthesis. Fruit seated
initially persisting,
and sometimes
tropical Asia,
but
has
to
extending
leaves
cupules.
a
off. Leaves
generally clustered, tripliveined
perulate.
About 100 species,
its dimerous flowers
characters
vegetative
although
help separate
the
along
In
sheath
a
at
species
of Neolitsea the bracts
species
in
genera: Neolitsea
usually
veined leaves and stalked umbels, these
shortshoots. Some Litsea
few
mostly
in other characters it
two
leaves and sessile umbels, these
the leafless parts of the innovations, while Litsea has
or
together
Australia.
recognized by
Litsea. Some
opposite), pinnately
as
to
several close
or
6, 4-locular; tepals equal,
small, plate-like cupule, the tepals sometimes
below. Terminal buds
usually clustered, tripliveined
along
on a
ultimately falling
glaucous
Neolitsea is best
is close
but
single
stamens
borne
alternate
(rarely
usually
also have rather
surrounding
frequently
in the axils of
large, cup-shaped
the terminalbuds
persist
the base of the leafless internodes.
19. Persea Mill
Inflorescence
paniculate-cymose
4-locular; tepals equal
of the fruit,
spreading
or
to
(type 2).
subequal,
reflexed,
a
cupule
pinnately veined, sometimes glaucous
genus with
regions
c.
200
species
of the Northern
in
tropical
Flowers trimerous,
half-erect
at
anthesis.
Tepals persistent
absent. Leaves alternate or
below. Terminal buds
Asia and America with
Hemisphere.
bisexual;
a
at
9,
the base
slightly clustered,
perulate.
few
stamens
A poorly defined
species
in
subtropical
H.
The
van
der Werff: An
above
description given
annotated
applies only
Asian Persea
and Phoebe and these
species
characters. In Asian Persea
species
in fruit, in Phoebe
persistent
are
the type
P. americana
species,
tepals
neither
tepals (the
and
partly
spreading
Mill.),
der Werff from
van
in fruit. A
equal tepals
others have
Surinam)
comprehensive study
tablishment of clear
spreading
to
on
fruit
reflexed
the fruit. Difficulties with
pressed against
has
deciduous in fruit
in
as
Phoebe),
strongly unequal tepals
unequal
or
in fruit while
3) persistent
(including
in fruit (but
equal tepals persistent
the fruit
pressed against
nor
and
persistent
due to the great variation found in American
3 much smaller than the inner
outer
(P. julianae
are
sometimes
are
usually separated
are
groups
are
which
species,
139
found floral differences between
not
two
tepals
in Persea. Some of these have
species placed
uous
they
delimitationin this group
generic
the
of Lauraceae
genera
the Asian
to
in the segregate genus Machilus. I have
placed
the
to the
key
species
one
which
decid-
are
of Persea s.l. and Phoebe is needed for the
es-
boundaries.
generic
20. Phoebe Nees
Inflorescence
paniculate-cymose
4-locular; tepals equal
and
Tropical Asia,
discussed
clasping
better
quite
21.
c.
sometimes
common
Phoebe
in the
in the FM
racemose or
pinnately veined,
in
can
be
9,
slightly clustered,
usually
not
from Persea
separated
It
fruiting stage.
than Persea,
area
with
not
a
that
is my
impression
perulate.
only
by
lack
the
that Phoebe is
with obovate leaves
species
species generally
few flower clusters
stamens
glaucous
vegetatively
perulate
are
buds.
near
the base of the inflorescence.
9, 4-locular; tepals equal,
accrescent
more or
less
erect at
floral tube. Leaves alternate
below. Terminal buds
not
A
perulate.
monotypic
very
stamens
similar
to
but the flowers
Eusideroxylon,
and the inflorescence is
basically
are
racemose
much
(laxly
Eusideroxylon).
Triadodaphne
Inflorescence
Kosterm.
paniculate (type 3).
tepals strongly unequal,
inner 3,
more or
veined,
not
only
or
stamens
Tepals persistent
from Borneo.
and with 9 fertile
paniculate
22.
only
is
Potoxylon
larger
anthesis.
absent. Leaves alternate
anthesis. Fruit enclosed in and fused with the
genus known
erect at
Kosterm.
Flowers trimerous, bisexual;
and
less
below. Terminal buds
in Phoebe and that Phoebe
Potoxylon
Inflorescence
cupule
glaucous
under Persea,
represented
a
Flowers trimerous, bisexual;
more or
species.
100
spreading tepals
vs.
(type 2).
subequal,
thebase of the fruit,
clasping
pinnately veined,
As
or
less
the
erect at
glaucous
outer
Flowers
3 much
anthesis. Fruit
trimerous, bisexual;
larger
not
below. Terminal buds
and almost
stamens
3, 2-locular;
completely covering
known. Leaves alternate and
not
perulate.
A
monotypic
the
pinnately
genus known
from Borneo.
Triadodaphne
nize it based
on
3 and the much
was
its
included by
Rohwer
strongly unequal tepals
deeper receptacle.
(1993 )
in Endiandra, but I
with the
Kostermans
outer
(1993)
3 much
prefer
larger
to
recog-
than the inner
included 2 additional
species
140
in
BLUMEA
had the
that
No.
46,
tepals
outer
species
1,
of T.
saw
greatly so)
initially placed.
was
I
specimen
shorter than the inner ones (but not
in Endiandra, where it
2001
Kosterm. from New Guinea and T.
inaequitepala (Kosterm.)
Kosterm. from the Solomon Islands. The
tepala
T.
T.
Triadodaphne,
—Vol.
I have
pachy-
inaequitepala
and I would include
not
seen
material of
pachytepala.
ACKNOWLEDGEMENTS
This
article is based
pleasant
1 and
2
largely
observations made
on
thanks to the Director
express my
and
one. I also thank Peter
productive
in
previously published
were
during
and Staff members
Stevens for
the Annals
4-week
a
who
at BO and
stay
did their
I would like
best to make
very
to
visit
my
a
critically reading the manuscript. Figures
of the Missouri
Botanical
Garden
83: 417
(1996).
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