Table Of Contentf
KEY TO THE GENERA OF
A Henk
van der
Werff^
LAURACEAE THE
IN
NEW
WORLDS
Abstract
The
A key to the twenty-nine genera of Lauraceae naturally occurring in the western hemisphere is presented.
A
key based on and vegetative characters only. brief statement on distribution and size of the genera is
floral
is
A
key
provided, together with a note on cupule shape. Spanish translation of the is included.
Lauraceae have, general, the reputation of both floral and fruit characters (Koslermans, 1957;
in
being After having studied the Hutchinson, 1964), or share a combination of these
difficult to identify.
neotropical members of this family for the last six problems. Keys that are based on floral and fruit
years, agree that the reputation well characters create special problems. As a rule, both
I fully is
deserved. There are various reasons for this. In floral and fruit characters are necessary for generic
the place, many collections are sterile or fruit- identification, but a single specimen rarely has both
first
need
ing and lack the floral characters needed for iden- flowers and fruits. Typically, fruits several
To date, there are no keys for the generic months for development, by which time the flowers
tification.
from an
identification of nonflowering specimens, and have long fallen Rarely, fruits earlier
it is off".
enough
unlikely such a key will become available in the flowering season persist long to be collected
may an such
specimens be recognized to with fresh flowers, but that exception. In
future. Sterile is
same
and do occur on
species by a botanist familiar with the family, but cases, flowers fruits not the
such instantaneous recognition does not occur, inflorescence. Occasionally, one finds flowers and
if
my
specimen remain even on the same inflorescence. In expe-
the likely to unidentified ''fruits''
is
and
genus and therefore inaccessible for a long time. rience, such "fruits'* are usually diseased de-
to
Fruiting specimens stand a better chance of being formed. Thus, because flowering specimens almost
never have key requires both flowers
because the cupule (or absence) con- fruits, a that
identified its
when and not workable and frustrate user.
useful information, especially old sta- fruits will its
tains is
am
mens remain on the cupule. However, not However, possible to construct a generic key
I it is
able to construct a workable key based on fruit for neotropical Lauraceae based solely on floral
and vegetative characters. From a taxonomic point and vegetative characters. I present such a key
American
specimens even though below, which includes genera.
of view, fruiting are valuable, all
whereas specimens are The most important floral characters used in
difficult to identify, sterile
number
of or no use. For botanists making forest this key, as well as in earlier keys, are of
little
stamens, number of anther per stamen, sexual
inventories or studying species diversity, sterile cells
voucher specimens are useful, hence collected, and condition of the flowers, and tepals equal or un-
end up on the desk of a specialist, where they often equal. These characters, although readily visible,
and
remain are often variable within a genus, this variation
unidentified.
Even with good flowering material at hand, iden- has rarely been incorporated in earlier keys. For
my
tification to genus often difficult. In opinion, example, Persea is usually presented as having
is
main reason problem no workable nine 4-celled stamens, whereas the neotropical spe-
the for this that
is
keys The published keys are partly obsolete cies can have nine or six stamens, these being all
exist.
some and some
(such as Mez, 1889), are of regional scope (Mac- 4-celled, all 2-celled, or 4- 2-celled.
1938; Kostermans, 1936; 1945), use Moreover, the tepals can be equal or unequal and
Bride, Allen,
W.
A. Gentry, M. Grayum, Rohwer, and D. Stevens critically read the manuscript and suggested improvement
' J.
W. Myers made
D. Stevens kindly translated the key into Spanish. the illustrations.
J.
Missouri Botanical Garden, P.O. Box 299, Louis, Missouri 63166-0299, U.S.A.
' St.
Ann. Missouri Box. Gard. 78: 377-387. 1991
378
Annals the
of
Garden
Missouri Botanical
deciduous or persistent in fruit. In order to accom- and II 4-celled, III 2-celled or sterile; whorls and
I
niodate these permutations of characters, at II 2-celled, whorl III sterile), but Endiicheria
it is
limes necessary to have a genus appear several anomnla has whorls T and II 2-celled and whorl
A
times in the key III 4-ceIIed. further variation that any of
is
While constructing key, have completely whorls and be (In some species
this I I, II, III sterile.
among
ignored any phylogenetic relationships the only whorl fertile, or only and II are fertile,
I is I
genera, mainly because phylogenetic schemes of or and are or only
II III fertile, III fertile;
is I
Lauraceae are partially based on fruit characters have not yet seen a species with only whorl II
(Kostermans, 1957) or are based on wood and Table presents the various androecial
fertile.)
1
among
bark anatomy (Richter, 1981). These supposedly combinations and their distributions neo-
phylogenetic classifications (which are quite differ- tropical Lauraceae, ''^Gamanthera'' excluded. If a
known
ent from each other) cannot readily be followed in key were to include combinations of
all
number number
the construction of a key based on characters. of stamens, their position and of
floral
my
Instead, aim to provide a workable key for anther cells in each whorl, one would end up with
is
How-
the identification of flowering Lauraceae. a very long and tedious (but workable) key.
Before presenting the key, I will briefly describe ever, I found that a shorter and more practical
made
the flower structure of Lauraceae. Examples are key can be by excluding the condition of the
Mez
based on American genera, and unusual conditions stamens of whorl as has been done by
III, first
of palaeotropical genera (such as dimerous flowers (1889). Exceptional species or groups of species,
Potameia) are not discussed. Typically, each such as the 2-celled species of Per sea and Carj-
in
flower has two whorls of three tepals each. The odaphnopsis and the Aiouea species with three
whorls are usually equal in size and shape, but stamens are keyed out separately.
there are exceptions. Tepals are unequal Following key, the distribution, approximate
in all tlie
Caryodaphnopsis^ many number and
species of in species of of species fruit type are briefly de-
When
Persea (Fig. Ic), in Anaueria, and in a few species scribed. an unusual leaf position occurs in
When
of Anlba and Licaria. tepals are unequal, a genus, this also mentioned.
is
A
the outer three are smaller, except in fAcaria, good magnifying glass, or preferably a dis-
where the outer three tepals can be larger (Fig. secting microscope, a necessity for identifying
is
unusual having Lauraceae.
Id). *'*'Chlorocardiu!n'' in the
is
tepals in whorls of four or irregular. The stamens Included in the key are several taxa which are
are arranged in four whorls of three each (with the not yet published {''''Chlorocardium,^^ ^^Gaman-
exception of ''''Chlororardium,'''' which has stamens thera,^^ and ^^Paraia''), Their inclusion in the key
in whorls of four) and are opposite the tepals. The absolutely not intended as their publication, but
is
whorls are counted from the outside to the center done to prevent the key from being obsolete at
is
Whorl moment
of the flower. opposite the outer tepals, the of publication.
I is
whorl II is facing the inner tepals, whorl III is
opposite whorl and whorl IV opposite whorl
I, II.
W
and
the exception of Litsea
ith ''''Cklorocar- Key New World
to Genera of Lauraceae
diuni^^^ whorl IV always staminodial or lacking.
is
Cassylha
Parasitic vine
leafless
1
The stamens of whorl nearly always have two
III
Shrubs or trees with green leaves 2
1
globose glands at their base, the exceptions being
2(1) '^Stamen" by complete fusion of 3 sta-
1,
Mezilanrus and Williamodendron which mens
these
in ''''Gamanthera''''
,
2 Stamens more than 3
glands are absent. In Phyllostemonodaphne and 1
Flowers 4
3(2) bisexual or staniinate
Urbanodendron stamens have
glands
at their
all
42
3 Flowers
pistillate
base. In Pleiirothyriiim the glands are strongly
may
enlarged and become completely
fused, sur- 4 Stamens more than 3 8
rounding stamens. In general, the stamens of 5(4) Stamens with 4 anther cells W^illiamode ndron
all
5 Stamens with 2 anther 6
cells
whorls and are (nearly) identical and seemingly
II
I
6(5) Leaves clustered near tips of branches; an-
The
form one whorl of stamens. exceptions are
six
ther cells extrorse; inflorescence a panicle,
Dicypellium and Phyllostemonodaphne, where
MezUaurus
racemose
ultimate divisions
the stamens of whorl have become tepaloid and 6 Leaves not clustered^ but alternate or op-
I
with only whorl and Stamens posite; position of anther cells variable (in
sterile, II III fertile.
L. cannella leaves somewhat clustered, but
of whorl occasionally from those of whorls
III differ
anther cells introrse); inflorescence panicu-
and number
of anther
II in cells. If diff'erent,
I cymose
liform, ultimate division 7
whorl III has usually fewer anther cells (whorls I 7(6) Only stamens of Whorl fertile, the cells
I
2
1
379
Volume Number 2 van der Werff
78,
New
Key World Lauraceae
1991 to
N', >.'..'
—
— &
Mez
Figure Flowers A. Nectandra grandiflora Nees {Gentry et ai 5976.3); B. Ocotea atirrensis
1. of:
—
Donn. Smith {Grayum 6868);— C. Persea Uebmannii Mez (Lshiki 1614); D. Licaria bracteata van der Werff
{Kunkel
56).
lateral and large, or lateral basal and small; 9(8) Flowers unisexual 1
inflorescences and flowers glabrous Aioiiea 9 Flowers bisexual 1
7 Only stamens of Whorl III fertile, the cells 10(9) Leaves deciduous in winter; flowers appear-
± Lindera
extrorse, introrse or apical, not lateral; ing before leaves
..„
inflorescences and/or flowers usually with 10 Plants evergreen, flowers and leaves present
same Endlicheria
amounts pubescence Licaria the time
varying of at
Outer 6 stamens each with 2 anther 9 11(9) Leaves opposite
8(4) cells „ 1
8 Outer 6 stamens each with 4 anther cells 23 11 Leaves alternate or clustered „ 15
m
49
380 Annals
of the
Garden
Missouri Botanical
Table Androecial configurations of American
1
Lau
raceae.
&
Whorls Caryodaphnopsis
pro
I, II, III 4-celled: parte,
C.innamomum
Chlororardiuniy pro Nec-
parte, Litscdy
tandrriy Oroten, Pnraia, Persea pro parte, Pleuroth-
Povcdodaph Rhodostemonodaph
yriu ne, nc^ Sas-
,
safras, Umhcllularia, Urhanodendron pro parte
&
Whorls Aiouea pro Aniba
I, II, III 2-celled: parte,
pro parte, Bcilsihmiedia, Caryodaphnopsis pro parte,
Cassythay Cryplocarya, Endlicheria pro parte, Ku-
blizkia pro parte, Lindera, Persea pro parte, IJrban-
odendron
pro parte
&
Cinnamomum
Whorls
pro
I II 4-ceIled, III 2-celled:
parte, Kubitzkia pro parte, Persea pro parte f.
&
Whorls
11 4-celled, III 0-celled: Persea pro parte
I
&
Whorls Endlicheria anomala
I II 2-celled, III 4-celled;
&
Whorls
Aiouea pro
II 2-celled, III 0-celled: parte,
I
AnaueriGj Aniba pro parte, Caryodaphnopsis pro parte
&
Whorl Aiouea
2-celled, II III 0-celled: pro parte
I
&
Whorl
I 0-celled, II III 4-celled; Dicypelliuni
& P
Whorl hy mono-
I 0-celled, II III 2.celled: Host e
daphne
&
Wliorl Williamodendron
II 0-ceIled, III 4-celled:
I
&
Wliorl II 0-celled, III 2-celled: Licariay Mezilaurus
I
12(11) Stamens 6, triangular, dark red, the fila-
ments wider than the anthers, united at the
hase; outer tepals about V^ as long as the
Anaueria
ones
inner
12 Stamens 6 or 9, filaments free, narrower
stamens never dark
than the anthers; red;
tepals equal or strongly unequal, outer ones >-
"'^
<14 ones 13
the size of inner
<V4
13(12) Outer tepals the size of inner tepals
....
Caryodaphnopsis
E
.__
3 Tepals equal
1 1
14(13) Floral tvihe deep, about as long as the tepals
Figure 2. Stamens (whorl or II) of:— A. Ncctandra
I
Cryptocarya —
..„ &
Mez (Kayap
reticuhita (R. P.) 161); B. Nectandra
much
14 Floral tube shallow, shorter than the —
niembranacea
(Sw.) Griseb. {Cuadros 2839); C. Ocotea
Beilschmiedia —
tepals
Mez
veraguensis (Meissner) (Stevens 6476); D. Ocotea
Whorl
stamens —
15(11) Filaments of of 111 united;
pulchella Mart. (Goetzke 210); E. Rhodostenwno-
Kubitzkia
flowers red
daphne
kunthiana Rohwer
(Nees) {Perry
s.n.).
15 Filaments flower color variable,
free; in-
some 16
cluding red species
in
Stamens Whorl
16(15) of sterile, tepaloid; fertile
I
stamens 6, representijig Whorls II and 111 short, much shorter than tepals; Andean
m
Phyllostemonodaphne 1,000-2,500 Persea
species, elev.
16 Stamens of Whorl fertile; fertile sta- 19 Staminodia (whorl IV) lacking; floral tube
I
mens 9 Whorl and Amazonian
or 6; 6, representing well developed; lowland species
if I
17 Aniba
canelilla or A. parviflora
II
17(16) Fertile stamens 9, each with 2 glands at the 20(18) Floral tube very shallow, scarcely visible at
Urhanodendron Beilschmiedia
base anthesis
17 stamens 6 or only those of Whorl 20 Floral tube well developed, about as long as
Fertile 9,
III with glands at their base 18 the tepals 21
18(17) Outer tepals ca. V2 as long as inner ones 21(20) Leaves with a fine, raised reticulation; floral
±
tube deep and narrow, the tepals spread-
(Fig. 1 c) 1
18 Tepals equal or nearly so (Fig. la and b) ing and flowers clearly wider than floral tube;
..
20 much
filaments narrower than anthers;
_
Cryptocarya
19(18) Staminodia (whorl IV) well developed, with staminodia well developed
sagittate or cordate apices; floral tube very 21 Leaves with lax reticulation or this not ap-
Volume Number 2 van der Werff 381
78,
New
1991 Key World Lauraceae
to
parent; floral tube shallow or deep; if deep, 34 Stamens of Whorl I fertile, not tepaloid 35
„.„.
tepals erect and flowers about as wide as 35(34) Outer tepals about Vi the size of the inner
36
tube; filaments frequently poorly ones
floral dif-
ferentiated from anthers; staniinodia present 35 Tepals equal or nearly so 37
or absent 22 36(35) Stamens of Whorls and II with a sterile,
I
_
22(21) Flowers (minutely) tomentellous, tomentose triangular leaf apex rounded or obtuse;
tip;
or with appressed pubescence, but never leaves whorled; Guyanas and adjacent Brazil
stamens and Venezuela Ocotea rubra
pruinose; filaments of usually densely
pubescent, poorly differentiated from an- 36 Stamens of Whorls and without a
I II sterile,
thers; leaves alternate or clustered Aniba triangular tip, the anther cells occupying the
22 Flowers usually glabrous, infrequently with entire anther; leaf apex various; leaves
al-
SE
scattered sometimes pruinose; ternate or whorled; widespread, from
hairs,
fila-
ments of stamens usually glabrous and U.S.A. to Brazil and Chile Per sea
dif-
ferentiated from anthers; leaves alternate 37(35) Staminal glands greatly enlarged, protruding
..
Aiouea between the outer 6 stamens and sometimes
23(8) Flowers unisexual 24 fused into a large, pillowlike mass; stamens
23 Flowers bisexual 27 of Whorls and with at least 2 lateral
I II
24(23) Young inflorescences enclosed decussate anther Pleurothyrium
in cells
bracts; flowers arranged in pseudo-umbels 37 Staminal glands not enlarged, not protruding
Lit sea between outer stamens; free; outer stamens
24 Young 38
inflorescences not enclosed in de- with introrse cells
cussate bracts; flowers arranged in panicles 38(37) Staminodia representing Whorl IV well de-
or racemes 25 veloped, with a cordate or sagittate apex;
25(24) Deciduous trees; flowers appearing before filaments of stamens as long as anthers or
or with young leaves, leaves often lobed longer 39
„„.
Sassafras 38 Staminodia representing Whorl IV small or
25 Evergreen plants with mature leaves when lacking; present, mostly without a cordate
if
flowering; leaves never lobed 26 or sagittate filaments of stamens shorter
tip;
26(25) Anther cells arranged in a low arch (Fig. than or as long as anthers (a few species of
anthers poorly from Ocotea have large staminodia with a cordate
2e); diff*erentiated
fil-
Rhodostemonodaphnc
aments but these have stamens with a very short
tip,
26 Anther arranged 2 rows 2c and filament) 40
cefls in (Fig.
d); anthers clearly diff"erentiated from 39(38) Leaves usually tripliveined, alternate, fre-
much
the narrower filament (rarely quently tufts of hairs present in the axils of
fila-
Cinnamomum
ments very short) Ocotca the lowermost veins
27(23) Flowers arranged in pseudo-umbel, this, when 39 Leaves pinnately veined, alternate or clus-
young, covered by bracts Umbellularia tered; axillary tufts of hairs lacking Persea
27 Flowers arranged in paniculiform inflores- 40(38) Leaves clustered; twigs with clusters of scars
cences, these never enclosed by bracts 28 from fallen bracts; tepals roundish, hyaline,
29
28(27) Leaves opposite with a pubescent base, otherwise glabrous
"Pa
28 Leaves alternate or whorled 31 ra a •ii
f
<
40
29(28) Outer tepals Vi the size of the inner ones; Leaves alternate; twigs without clusters of
from
leaves often strongly triveined scars faflen bracts; tepals not roundish,
Caryodaphnopsis
not hyaline, glabrous or with
difi'erent dis-
29 Tepals equal; leaves pinnately veined 30 tribution of pubescence 41
30(29) Stamens 12 or more; leaves glabrous or 4 (40) Anther cells arranged in 2 vertical rows (Fig.
1
nearly so on lower surface ''''Chlorocardiuin^ 2c and stamens and inner face of tepals
d);
30 Stamens leaves ferruginous tomentose or glabrous or variously pubescent, rarely pa-
9;
tomentellous on lower surface piflose papillose, the anther cells clearly
(if
_
Nectandra
oppositifoUa in 2 vertical rows); te[)als free at base, falling
31(28) Anther cells on the flat tip of columnar sta- individually in old flowers; a few old stamens
mens Povedadaphne sometimes present on cupule of young
fruits;
31 Anther cells not apical; stamens not colum- tepals at anthesis erect (Fig. lb) or spreading
32 Ocotea
nar „
32(31) All stamens with 2 glands at their base, the 41 Anther cells arranged in an arc (Fig. 2a and
glands free Urbanodendron b); stamens and inner face of tepals papil-
32 Only stamens of Whorl III with glands (in lose; tepals united at the base, usually falling
Pleurothyrium glands greatly enlarged and as a unit (together with stamens) in old flow-
sometimes fused) 33 ers; stamens rarely present on cupule of
Stamens Whorl young
33(32) of III with fused filaments; fruits; tepals at anthesis spreading
flowers red Kubilzkla (Fig. la) Nectandra
33 Stamens of Whorl III with free filaments; if 42(3) Plants cold-season deciduous and/or inflo-
filaments seemingly fused some Ocotea rescences subumbellate and young flowers
(in
43
species), then flowers white to yellow, never enclosed in involucrate bracts
42
red 34 Plants evergreen and inflorescences panic-
Stamens Whorl
34(33) of devoid of anther ulate, the flowers not enclosed involucrate
I cells, in
m
tepaloid DicypeUiu bracts 45
2 46
1
382 Annals of the
Garden
Missouri Botanical
43(42) Inflorescences racemose or paniculate; leaves 12 Estambres 6 o 9, nunca rojo-obscuros, con
mas delgados que anteras
often trilobed Sassafras filamentos las y
muy
con
43 Inflorescences subunibeflate; leaves never libres; tepalos iguales o desiguales
menos
trilobed 44 los exteriores !4 o del largo de los
13
44(43) Plants deciduous; pseudo-umbels sessile or interiores
menos
nearly so Lindera 13(12) Tepalos exteriores 14 o del largo de
Caryodaphnopsis
44 pseudo-um-
Plants deciduous or evergreen; los interiores
pedunculate Litsea 3 Tepalos iguales
bels 1 _ 1
Tubo como
45(42) Staminodes straplike, the filaments about as 14(13) floral profundo, tan largo los
Cryptocarya
wide as the anther; traces of four on tepalos
locelli
mucho mas
Rhoilostemonodaphne 4 Tubo poco profundo, corto
each anther floral
1
Beilschmiedia
45 Staminodes club-shaped, the filaments nar- que los tepalos
rower than the anthers 46 15(11) Estambres del Verticilo III con filamentos
Kuhitzkia
46(45) Anthers with 4 remnants of leaves unidos; flores rojas
locelli;
Ocolea 5 Estambres con filamentos libres; flores rara-
alternate
_ 1
46 Anthers with 2 remnants of leaves mente rojas
locelli; 1
whorled Endlichcria 16(15) Estambres del Verticilo esteriles, tepa-
alternate or I
estambres representando
loides; fertiles 6,
Phyllostenwnodaphne
los Verticilos II y III
16 Estambres estambres
Clave Para los Generos de del Verticilo I fertiles;
9 6 6 entonces representando
Mundo fertiles 6, si
Nuevo
Lauraceae
del
17
los Verticilos I y II
Trepadoras parasitas y afilas CassyUia 17(16) Estambres fertiles 9, cada uno con 2 glan-
1
Urbanodendron
Arbustos o arboles con hojas verdes 2 dulas en base
1 la
2(1) Estambre aparentemente 1 por fusion com- 7 Estambres fertiles 6 6 9, solo los del Verticilo
1
pleta de 3 estambres ^^Gamanthera'*^ con glandulas en base 18
III la
Estambres claramente mas de 3 18(17) Tepalos exteriores cerca de del largo de
1 ^/z
3(2) Flores bisexuales o estaminadas 4 los interiores (Fig. Ic) 19
_
42
3 Flores pistiladas 18 Tepalos iguales o casi iguales (Fig. la y b)
20
Estambres 3 5
4(3)
_
mas
4 Estambres de 3 8 19(18) Estaminodios bien desarrollados, con apices
muy
5(4) Anteras con 4 tecas Williamodendron sagitados o cordados; tubo floral corto,
5 Anteras con 2 tecas 6 mucho mas corto que los tepalos; especies
msnm
6(5) Hojas agrupadas en los extremus de las ra- andinas de 1,000-2,500 Persea
„.
mas; tecas extrorsas; inflorescencia una pa- 19 Estaminodios ausentes; tubo bien de-
floral
nicula con las divisiones ultimas racemosas sarrollados; especies amazonicas de eleva-
Mezilaurus ciones hajas Aniha ranelilla o A. parvijlora
6 Hojas alternas u opuestas, no agrupadas; 20(18) Tubo poco profundo, escasamente
floral vis-
orientacion de las tecas variable (en L. can- en antesis Beilschmiedia
ible la
mas menos
nella hojas algo agrupadas, pero entonces 20 Tubo bien desarrollado, o
floral
mismo que 21
las tecas introrsas); inflorescencia paniculi- del largo los tepalos
forme, divisiones ultimas cimosas 7 21(20) Hojas con una reticulacion fina promi-
y
7(6) Solamente los estambres del Verticilo I fer- nente; tubo floral profundo y angosto, te-
± Umbo mas
tiles, tecas laterales y grandes o lateral-ba- palos patentes y el claramente
mucho mas
sales petjuenas; inflorescencias y flores ancho que tubo; filamentos
y el
glabras Aiouea angostos que anteras; estaminodios bien
las
Solamente estambres Cryptocarya
7 los del V^erticilo III desarrollados
tecas extrorsas, introrsas o inapica 21 Hojas con una reticulacion laxa o inconspi-
fertiles,
y/o
nunca laterales; inflorescencias cua; tubo poco profundo o profundo,
les, flo- floral
res usualmente pubescentes Licaria profundo entonces los tepalos erectos
si y
8(4) Anteras de los 6 estambres exleriores con limbo tan ancho como el tubo; filamentos
el
2 tecas 9 frecuentemente poco diferenciados de las
8 Anteras de los 6 estambres exteriores con anteras; estaminodios presentes o ausentes
23 22
4
tecas
menudamente
9(8) Flores unisexuales 10 22(21) Flores tornentosas o aplicado-
9 Flores Disexuaies pubescentes, nunca prulnosas; filamentos con
hi 1
1
10(9) Hojas deciduas en invierno, flores produci- frecuencia densamente [)ubescentes, poco
el
das antes que las hojas Lindera diferenciados de las anteras; hojas alternas
Aniba
10 Hojas no deciduas, hojas presentes o agrupadas
flores y
a vez Endlicheria 22 Flores usualmente glabras, infrecuente-
la
11(9) Hojas opuestas mente con tricomas dispersas, a veces prui-
1
11 Hojas alternas o agrupadas 15 nosas; filamentos usualmente glabros y di-
12(11) Estambres rojo-obscuros, con filamentos ferenciados de anteras; hojas alternas
6, las
mas anchos que anteras unidos en Aiouea
las y la
24
base; tepalos exteriores cerca de V2 del largo 23(8) Flores unisexuales
de los interiores Anaueria 23 Flores bisexuales 2 7
Volume Number
2 van der Werff 383
78,
New
1991 Key World Lauraceae
to
24(23) Inflorescencias jovenes envueltas en brac- pujando entre los 6 estambres exteriores
y
teas decusadas; flores en pseudo-umbelas a veces fusionadas en una masa grande como
...
Litsea almohada; estambres de
los Verticilos y
II
I
24 Inflorescencias jovenes no envueltas en con al menos 2 de las tecas laterales
m
bracteas decusadas; flores en paniculas o Plea rot hyriu
racimos 25 37 Glandulas estaminales no agrandadas
ni fu-
25(24) Arboles deciduos, flores producidas antes o sionadas; estambres exteriores con todas
las
junto con las hojas; hojas frecuentemente tecas introrsas 38
lobadas Sassafras 38(37) Estaminodios representando Verticilo IV
el
25
Arboles o arbustos perennifolios, flores prod- bien desarroUados, con apices cordados o
ucidas con las hojas maduras; hojas nunca sagitados; filamentos iguales o mas largos
lobadas 26 que anteras 39
las
„
26(25) Tecas dispuestas en un arco bajo; anteras 38 Estaminodios representando Verticilo IV au-
poco diferenciadas de los filamentos (Fig. sentes o presentes pequenos y mayor-
si
Rhodostemonodaphne
2e) mente sin apices cordados o sagitados;
fi-
..
26 Tecas dispuestas en 2 hileras; anteras clara- lamentos iguales o mas cortos que las anteras
mas
mente anchas que los filamentos (rara- (unas pocas especies de Ocotea tienen es-
mente
filamentos niuy cortos) (Fig. 2c y d) taminodios grandes con apices cordados, pero
muy
Ocotea 40
estas tienen filamentos cortos)
27(23) Flores en pseudo-umbelas, estas cubiertas 39(38) Hojas usualmente trinervias, alternas, fre-
por bracteas cuando jovenes Umbellularia cuentemente con fasciculos de tricomas pre-
27 Flores en inflorescencias paniculiformes, es- sentes en las axilas de los nervios basales
.,
tas nunca envueltas por bracteas 28 Cinnamornum
28(27) Hojas opuestas 29 39 Hojas pinnatinervias, alternas o agrupadas,
28 Hojas alternas o verticiladas 31 sin fasciculos de tricomas axilares Persea
29(28) Tepalos exteriores del tamafio de los in- 40(38) Hojas agrupadas; ramitas con fasciculos de
!/4
mas
teriores o pequenos; hojas con frecuen- cicatrices de bracteas caidas; tepalos algo
cia fuertemente trinervias Caryodaphnopsis redondos, hialinos, pubescentes solamente
29 Tepalos iguales; hojas pinnatinervias 30 en base "Pa raia i»
la
30(29) Estambres 12 o mas; hojas glabras o casi 40 Hojas alternas; ramitas sin fasciculos de
ci-
glabras en el enves ''''Chlorocardium'''* catrices; tepalos ni redondos ni hialinos, gla-
30
Estambres 9; hojas ferrugineo-tomentosas bros o con la pubescencia distribuida de otra
en enves Nectandra oppositifolia forma 41
el
31(28) Tecas en el apice piano de los estambres 41(40) Tecas dispuestas en 2 hileras (Fig. 2c y d);
columnares Povedadaphne estambres caras adaxiales de tepalos
y los
31 Tecas no apicales, estambres no columnares glabros o variadamente pubescentes, rara-
32 mente
papilosos papilosos, las tecas clara-
(si
._
Todo
32(31) los estambres con 2 glandulas en la mente en 2 hileras); tepalos libres en la base
base, las glandulas libres Urbanodendron y cayendose individualmente; unos pocos
32 Solamente los estambres del Verticilo III con estambres viejos frecuentemente presentes
glandulas (en Pleurothyrium las glandulas en la cupula de los frutos jovenes; tepalos
muy
agrandadas y a veces fusionadas) 33 erectos o patentes en la antesis (Fig. 2b)
...
33(32) Estambres del Verticilo III con filamentos Ocotea
fusionados; flores rojas Kubitzkia 41 Tecas dispuestas en un arco (Fig. 2a y b);
33 Estambres del Verticilo III con filamentos estambres y caras adaxiales de los tepalos
o aparentemente fusionados (en papilosas; tepalos unidos apenas en base,
libres, si la
algunas especies de Ocotea)^ entonces usualmente cayendose juntos juntos con
las
(y
flores blancas a amarillas, nunca rojas 34 los estambres); estambres raramente pre-
34(33) Estambres del Verticilo I esteriles y tepa- sentes en la cupula de los frutos jovenes;
Dicypellium en
loides tepalos patentes antesis (Fig. la)
la
34 Estambres no Nectandra
del Verticilo I fertiles y te-
paloides 35 42(3) Plantas deciduas en invierno y/o
el inflo-
35(34) Tepalos exteriores cerca de ^A del tamano rescencias subumbeladas y flores jovenes en-
de los interiores 36 vueltas por bracteas involucrales 43
35 Tepalos iguales o casi iguales 37 42 Plantas perennifolias con inflorescencias
36(35) Estambres de los Verticilos y II con apices paniculadas, flores no envueltas por bracteas
1
triangulares y esteriles; hojas con apices re- involucrales 45
dondeados u obtusos; Guayanas y areas ad- 43(42) Inflorescencias racemosas o paniculadas; ho-
yacentes de Brasil y Venezuela Ocotea rubra jas frecuentemente trilobadas Sassafras
__.
36 43
Estambres de Verticilos I y II sin apices Inflorescencias subumbeladas; hojas nunca
ocupando 44
triangulares y esteriles, las tecas trilobadas
toda la antera; hojas raramente con apices 44(43) Plantas deciduas; pseudo-umbelas sesiles o
redondeados u obtusos; ampliamente Lindera
distri- casi sesiles
44
buida desde el sur de los Estados Unidos Plantas deciduas o perennifolias; pseudo-
hasta Brasil Chile Persea umbelas pedunculadas Litsea
y
muy
37(35) Glandulas estaminales agrandadas, em- 45(42) Estaminodios ligulados, filamentos igual de
384 Annals of the
Garden
Missouri Botanical
ancho que las anteras, cada antera con ves- ed on a naked pedicel. It occurs in lowland forests
Rhodostcmonodaphne
tigios de 4 tecas from Costa Rica to Peru and Brazil. The seven
mas
45 Estaminodios claviformes, filamentos
occur Indochina, with one
palaeotropical species in
46
delgados que anteras
las
New
Guinea (Kostermans,
extending
species to
46(45) Anteras de estaminodios con vesligios de
los
1974).
Ocotea
4 tecas; hojas alternas
46 Anteras de estaminodios con vestigios de
los
Cassytha
L.
2 tecas; hojas alternas o verticiladas
Endlicheria
One from Mexico
species the Neotropics, to
in
Aiouea often found near the coast, but also locally
Aublet Brazil,
common More
Mauritia swamps. species occur
in
Probably a polyph,yl1etic genus in the currently c 4r u u
.l t-u
. . ,. andi South • Ihe genus has been
Australia Africa.
•^ in
,1
accepted sense, as has been noted by several1 au- •
u
^ •. own r r
' : ipli aiceidi in its ta.mij-li y-be^cauuste ot its parasitic
1988: Rohwer van
thors (Burner, et in press;
al.,
^ ^ habit, but hi ere retained in the Lauraceae.
iS
r^,
der Werff, 1987b, 1988).V The genus ,has ,been
ty
monographed by Renner (1982) and consists of '^Chlorocardium
about 20 species, raneine from southern Mexico n
r j
A
. genus wi.t,h ^two species, one trom Lruyana and
^
to southern rB^razil1 andI Paraguay. MostIspIeciI.es have a a^^ ^^a
S^^uri• nam, the ot.hu er ftrom Aa mazonian th?cuador and
„
, , n
the seated on a shallow cupule with a thickened
fruit
^ ,. --
adjacent, Coli ombiia• , previ• ousli y i• nciludi edi i• n iJcot# ea,
pedicel.
wood, and
but differing in characters of flowers^
The Ecuadorian/Colombian
species
Anaiieria Kostermans leaf position.
incompletely known. Fruits are described as large
is
A known Amazonian
monotyyp^ic^^genus from
for- ^
/, -77 X 5r cm)\, wi-t^hu a normal1 or very 1lar„ge ^cup^ul1e^.
(to
and The
near the border of Brazil Peru. seeds
ests
Cuinamomum
A
Schaeffer
cupule
are said to be edible after roasting. is
lacking Flowers have green tcpals and dark
in fruit.
A
poorly understood genus occurring in
large,
stamens, very unusual Lauraceae.
red in
Asia, Australia, and the Americas. Possibly with
known
Aniba Aublet more than 50 species in the Neotropics,
.11
West
from Mexico and the Indies to southern Brazil
™
Aniba
Recently revised by Kubitzki (1982). in-
^^^, tP>araguay. tT.h! e r -. • suub»tendj ed1 iby a smal1l1
fruit
•^ is
•'
40-50 many
eludes species, with pleasantly aro- .
. i^^^^io
^ ^ cupule, t.hi. s of^^ten crowned, wi• t,h persi•st, e.nt tepals.
'
1. r
matic yellow wood (rose.wo.od. 1 • disti1l1led1 from
oil is
,,,. ^ c^ ^..^^a ;^
y Ihe neotropi.cal, speci. es ,have otten \been ,treated in
^
Aniba from Costa
rosaeodora). distributed
It is Cinnamomum.
Phoebe, seem placed
but better in
Rica southern Brazil and Bolivia. Most species ,, r
to
Many A. si. an speci.es, i• ncl\ uad*ing„ .v,e^r.u,,n«i^, .w.n.\i.c\^nU
^ (..
occur the lowlands, but has also been collected
in
it
cinnamon, have opposite leaves, but neo-
yields all
Andes up 1,800 m. The ,1s1eated
in the to fruit is r ^-^^a
r ' ^ . , speci.es .have al1.ternat.e, most*li y *tnpliveined
tropical
a rather deep cupule, which often lenticellate ^.
in is u
^ ^ l, eaves. Cinnamomum, as accept* edj here, • possibuily
is
.'
have
or warty. Several species clustered leaves.
polyphyletic.
Beilschmiedia Nees
Brown
Cryptocarya
R.
In the Neotropics a poorly understood genus A poorly understood pantropical genus,
large,
10-20 Kostermans
by
with species, last revised ^^^ ^^ fewer neotropical species, mostly in
^j^j^
(1938), who accepted 15 species. It has been col- southern Brazil and Chile, but also known from
lected from Mexico to southern Brazil and in the French Guyana and adjacent Andean Ven-
Brazil,
West
Ecuador, and P The almost
ezuela, fruit
is
Heilschmiedia equally poorly un-
laeotropics, is completely enclosed the cupule, which has only
in
many
by more
derstood, but represented species. ^
is small, apical pore.
&
Caryodaphnopsis Shaw DirypeUium Nees Martius
Airy
A
Only recently reported from the Neotropics (van small genus of two rarely collected species.
&
der Werff Richter, 1985), this genus includes restricted lo Amazonian Brazil, reviewed by Roh-
published neotropical species, with several more wer (1988). The cupule cupshaped, and has a
five is
The
awaiting publication. The strongly unequal tepals double margin. tepals are persistent in fruit,
One Dicypellium caryophyllaceum, has
and opposite leaves are diagnostic. Fruits (in neo- species,
tropical species) are round or pearshaped, and seat- been heavily exploited because of its aromatic bark.
Volume Number 2 van der Werff 385
78.
New
1991 Key World Lauraceae
to
Endlicheria Nees shaped cupule. Leaves of Litsea glaucescens are
bay
used as a spice, similar to leaves.
A 40
neotropical genus of about species, re-
ported from Costa Rica, the Lesser Antilles south Mezilaurus Taubert
Paraguay, and southern The genus
to Brazil.
is
A
neotropical genus of 16 species occurring
in
(Rohwer The
cupule
polyphyletic et in press).
al.,
Amazon
the basin (including Bolivia, Peru, Colom-
most have and
variable; species a thick, fleshy
is
Venezuela, and the three Guyanas), revised
bia,
smooth cupule. Several species have whorled leaves.
by van der Werff (1987a). The seated on
fruit
is
Gamanthera^ The
'''' a small, platelike cupule. flowers are very
two genera
similar to those of Licaria, but the
A
monotypic genus recently discovered Costa
in
differ in cupule shape, leaf position, inflorescence
Rica, characterized by the presence of only one
and wood anatomy. have
type All species leaves
"stamen," synandrium formed by com-
actually a
Two
clustered at tips of branches. species with 4-
stamens comm.).
plete fusion of three (Endress, pers.
were
celled anthers recently transferred to Wil-
The number synandrium can be
of of the
locelli 3,
&
liamodcadron (Kubitzki Richter, 1987).
The
or cupule rather deeply cupshaped.
2, 1. is
margin and Nectaadra
with a double persistent, reflexed te- Rol. ex Rottb.
The
monoecious.
only species
pals.
is
A
large, neotropical genus of about 120 species,
currently under revision by Rohwer. occurs
J. It
Kubitzkia van der Werff*
from southern Florida and Mexico to Argentina.
A
genus
neotropical of one, possibly two, species. Fruits are seated in a cupshaped (sometimes small)
known
from Guyana, Surinam, Venezuela, and
cupule. Tepals are spreading at anthesis.
Rohwer who
Brazil, reviewed by (1988), accepts
Systemonodaphne name Ocotea
as the valid for this genus. Aublet
The cupule cupshaped, with a double margin
is
The
largest genus of Lauraceae in the Neotrop-
and Arguments
persistent, reflexed tepals. for re-
300 known
from
with at least species. also
ics, It
is
name Systemonodaphne
jecting the are discussed
Madagascar and The genus very
tropical Africa.
is
by van der Werff' (1986).
and dumping ground
variable serves as a for species
accommodated
that cannot be readily other gen-
in
Licaria Aublet
era. Its distribution in the Neotropics from Mex-
is
A
40
neotropical genus of about species, revised ico and southern Florida to Argentina. Shape and
by Kurz (1983). occurs from southern Florida of cupule variable and ranges from small
It size
is
and Mexico to southern Brazil and BoUvia. The and platelike to cupshaped, sometimes with a dou-
cupule cupshaped and has a double margin; ble margin or with persistent tepals. Tepals are
is
however, in some species the double margin is erect or spreading at anthesis.
A
scarcely noticeable. few species have opposite
leaves. ''^Paraia''
A
monotypic genus known from Amazonian
Bra-
Lindern Thunberg
The cupule has a double margin with persistent
zil.
Lindera is represented by three species in east- tepals. Tepals are erect at anthesis.
ern North America and an hundred
additional in
Persea
The Miller
on
Asia. fruit seated a small, platelike cupule.
is
Two
of the American species are rare; one was
An
incompletely understood genus with
at least
only recently described (Wofford, 1983). Twigs,
80
neotropical species, but better represented in
common
and American
leaves, fruits of the species
and one Canary
Asia with species in the Islands.
are used prepare a fragrant
to tea.
The fruit can be seated on a naked pedicel (as in
Persea aniericana, the type species) or sub-
is
Lamarck
Litsea
tended by persistent, indurate tepals (as in the great
New World A A
In the Litsea represented by about majority of species). cupule never present.
is is
known
five species, from the U.S.A., the mountains worldwide survey needed to determine Machi-
is if
of Mexico, and Costa Rica. Most species (several /a5 belongs in Persea (as accepted here) or should
is
The
hundred) occur in Asia south to Australia and the be recognized as a distinct genus. neotropical
Pacific Islands. The fruit subtended by a cup- species have been revised by Kopp(1966). Persea
is
386 Annals of the
Garden
Missouri Botanical
Mez
americana Urbanodendron
is widely cultivated for its edible fruit.
A
few neotropical species have clustered leaves.
A
neotropical genus of three species, restricted
The known two
Phoebe Nees southern cupules are
to Brazil. in
more
species: double-rimmed, with or less persis-
Phoebe
here considered a palaeotropical ge-
is
The genus has been revised by Rohwer
tent tepals.
Phoebe
nus. Neotropical species formerly placed in
(1988).
Cinnamomurn;
are, for the larger part, included in
&
a few belong to Ocotea. Williamodendron Kubitzki Richter
A
Ph yUostenwnodaphne Kostermans genus two
neotropical with species, infre-
known
quently collected, but from Costa Rica,
An
monotypic genus from
infrequently collected,
northern Colombia, Amazonia, and southern Brazil.
The
southern
the Atlantic rainforests in Brazil. fruit
The number more
of species increase as
will likpely
double-rimmed The genus
seated a cupule.
in
is
become The cupule small
collections available.
is
Rohwer
was revised by (1988).
and platelike, resembling that of Mezilaurus. Leaves
Pleurothyrium Nees are clustered at the tips of branches.
A 40
neotropical genus with close species,
to
Literature Cited
known from Guatemala south to Peru, Bolivia, and
A
Allen, C. K. 1945. Studies in Lauraceae VI pre-
The
seated a rather deep, cup-
Brazil. fruit in
is
Mexican and Amer-
liminary overview of the (.entral
n
f
s,haped, cupul1e, wh1 i- ch1 • often warty or l1 enti• cellate.
us
280-434.
.^^^ ^^^^^^ ^^^^^ 26:
j ^^^^^,^j
have pronoimced marginal
Several species a vein, Burger, W. 1988. A new genus Lauraceae from
C. of
an unusual feature in Lauraceae, or clustered leaves. Costa Rica, with comments on problems of generic
The genus heing revised by van der WerfT. and specific delimitation within the family. Brittonia
is
275-282.
40:
Povedadaphne Burger Hutchinson, 1964. The Genera of Flowering Plants,
J.
Volume Clarendon Oxford.
Press,
I.
A
monotypic genus apparently Cos- A
restricted to Kopp, 1966. taxonomic revision of the genus
L.
The Persea Western Hemisphere. Mem. New York
ta Rica. rather large fruit subtended by a in the
is
11
Bot. Card. 14: 20.
The
small, platelike cupule. fruiting pedicel swol-
is
Kostermans, A. G. H. 1936. Lauraceae. Flora
J. /;/.
columnar stamens,
len. Distinctive are the nine
of Suriname. Meded. Kolon. Amsterdam, Afd.
Inst.
each
with four apical
locelli. Handelsmus. Volume 244-337.
11.
1938. Revision of the Lauraceae V. Rec.
.
&
R
hodostenionoddphne Rohwer Kubitzki 834-931.
Trav. Bot. Neerl. 35:
Conmiun.
1957. Lauraceae. Forest Res. Inst.
.
A
known
poorly geims of about a dozen species,
1-64.
57:
common, A
two of which are rather while several 1974. monograph of Caryodaphnopsis A.
.
123-
The Shaw. Reinwardtia 137.
need genus has 9:
species are of description.
in
*]f r-^D- JO
*r>
Kubitzki, K. 1982. Anlba. Flora Neotropica 31:
In:
Ibeen reported from Costa Kica -i and reru. ^
to Brazil
^^
1-84.
^
Vhe cupule deeply cupshaped and rather large. &
is Williamodendron
H. G. RiCHTER. 1987.
&
Kubitzki Richter, a new genus of neotropical Lau-
Sassafras Presl
49-58.
raceae. Bot. Jahrb. Syst. 109:
Kurz, H. 1983. Fortpflanzungsbiologie einiger Gattun-
A
genus one widespread
small with three species,
gen neotropischer Lauraceen und Revision der Gat-
in the eastern U.S.A., the other two in China,
tung Licaria (Lauraceae). Ph.D. Thesis, Univ. of
The
Leaves are frequently lobed. cupule cup- Hamburg, Hamburg.
is
MacBride, 1938. Lauraceae. Flora of Peru. Field
shaped, often crowned with remnants of the tepals. F. In:
819-931.
Mus.
Nat. Hist. Bot. Ser. 13(2):
Bark and root have been used to prepare tea or
Mez, 1889. Lauraceae Americanae. Jahrb. Konigl.
C.
root beer.
1-556.
Bot. Cart. Berlin 5:
Renner, 1982. Aiouea. Flora Neotropica 31:
S. In:
System onodaphne
85
116.
Richter, H. G. 1981. Anatomic des sekundaren Xy-
See Kubitzkia.
lems und der Rinde der Lauraceae. Sonderb. Natur-
Hamburg 1-148.
Vereins
wiss. 5:
VmbeUalaria Nut
tall
Rohwer, G. 1988. The genera Dicypellium, Phyl-
J.
lostenwnodaphney Systemonodaphne and Urbano-
A
monotypic genus occurring California and
in
157-
drndron 110:
(Lauraceae). Bot. Jahrb. Syst.
southern Oregon The subtended
(U.S.A.).
fruit is —
171.
The
by a small, platelike cupule. leaves are used &
H. G. Richter H. van der Werff. 1991.
,
as a spice; the wood valued tor cabinetmaking. Two new genera of neotropical Lauraceae and critical
is