Phylogenetic Considerations of Euphorbiaceae Tribe Aleuritideae

10 Pages·1999·8.1 MB·

by  Peter C Van Welzen

PHYLOGENETIC
van
Peter Welzen*
C.
Wolfgang Stuppy-
CONSIDERATIONS OF
EUPHORBIACEAE
TRIBE
ALEURITIDEAE
1
between the genera, though small, an- based
M„i Ueuntideae.
Ci ,,„-, ,,-). lor On.' s.-leelcd
/ till,,- <il tin- ..
I
was tentatively supported lor only subtribe iarciinae (moiiogeneriel. irolonogv nmac
Iv
( (
geneia). (.i.wM-nna.' genera), prohahlv paraphy
{2 - miIiImI..-. !?
I'll.- 1 . 1 1 ti < ' 1 i- let ic.
synonymy
ently placed in of Tunnodia by Radcliffe-Smith) and Tunnodia group with subtribe
Nowadays
siiblribc Meiinlinae comprises three with small glandular bans: however, they differ
in
small genera: Aleurites (2 species), Reutealis (1 spe- leaf venation, presence of a caruncle, and persis-
cies), and Vernicia (3 species). Midler (1866) had tence of stipules. \ln. an tsubtribe
\I<ii>ni<>ph\t»r>
former iimlcd die three genera within broader Crotonogyninael
l\ a -tin- a -J d by its vining hab-
is. -I
.
Shaw (VXu)
Aleurites. Airy subdivided Aleurites s.l. palmate venation, and stinging hairs; the other
it,
two genera, Grotonogyne and Cyrtogamme, are
zingia, and Tapoidcs, thought to In closelv related quite similar and -bare thai aetei such as dioecy
s
to Aleurites sd. Uecentlv. Tapaiiles was placed in and fused petals. The inauilv Mriean Crosserinae
Web-
siil.inbc irosserinae Ueuritideae)
(tribe b\ die largest subtribe. with seven genera,
( is all di-
He
(1994b). considered Deutzianthus
ster (with oecious and with free petals. The morphological
hierzingia as synonym) belong among
lo to tribe Jatro- variability these genera great and their
is
pheae and
not tribe Aleuritideae.
distributions (Table disparate (South America,
1)
Ma-
Revision of subtribe Aleuritinae Flora
for Southeast perhaps
\ln. a. \sia). indi. -alive ol a par-
ma
suggest- differences between genera
the
les/, to aphy group.
lelie
be narrow and reasonable
reticulate. therefore
is
Il
ask whether
to this taxnriniuic di\rsion instilled
is
Menu
and whether the genera and
split siiblribc
The
tinae are monophylelic. phy logenetic analysis
herein for genera of tribe Aleuritideae (Table
all
1)
The
will address these questions. characters used in the cladistic analysis are
explained Appendix
For tribe Aleuritideae, the four subtribes Garci- in 1; the data matrix is pre-
The
sented in Table 2. character states were ob-
inae show various degrees homogeneity (see Ta- tained from herbarium specimens (Appendix '> and
,,l
American
ble for classification). suhtrihe literature (Table
1 1).
m
Garciinae moiiogenenc and almost by definition following characters were not included
is
I'll.-
homogenous; however, the two species of Garcia the cladistic analysis because they are typical
for
number
differ considerably in the stain. n |_"> ver- single taxa and. therefore, are presumably autapo-
63 The
sus to 160). two genera of subtribe Neo- morphies: whorled leaves; ovary 2-loculai „'!><<>
ill,
boulorunae share the ahs.-iice of petals and leaves ileiulnmY. amplexieanl stipules; bracts forming
a
Nnppy
flic
KM.
IJISII. ltd. an.l l.'a.e ll.ai.kcd for Use' ol"
id three e<lilors provided very useful comments.
- Hijksherbariiirn/Hortus llotaniciis. Diversity of Leiden. P.O. Box <)ol4. 2300
I
I
-
IVibc M.Mirili
Suhtribe
Al
Mm
spec.es, Mdles.a, \ustrali
rites
Stuppy
et (1999)
al.
species, China, Indochina Stuppy et (1999)
al.
(1<H5)
l.un.lell
Ducke
\,„<.m,>hc„h\ (1934)
&
Elffers Taylor (1956)
Leandri (1940)
&
Pax Hoffmann
(1912)
Lanjouw
(1932)
&
Pax Hoffmann
(1912)
Shaw
Airy (1971)
&
Pax Hoffmann
(1912)
&
Pax Hoffmann
(1912)
&
Pax Hoffmann
(1912)
Radcliffe-Smith (1988)
&
Pax Hoffmann
(1914)
" '
I'lll r:»l.:ii.
(
Sum
2 species. Indochina.
cnne •a), al abaual id.unU pre-
\( ,:i,t, -.uliiii. ii,-ii,
sligmas
sent, crater-like; hiiid to inultilid {(',n>h>n
n^\nr): stammate receptacle wilh disc-like glands
Alei
now
(Cyrtogonone); pilose receptacle (Domohinea,
synonym
a of 'faiinorfin. see Uad< lific-Sinith. 1998):
&
may
uisite (Nixon Carpenter, 1993), but facilitate
ossera): lian.
-
^lamls surface of leaves (Manniophytim); cliara- [• polan/; tini.. I>r>,lz;,i"lh -is. roosidet « •: in
i 1
il
(
Shaw
Airy (19671 to be a close relative of Aleurites,
seeds carunculate (Neoboutonia);
probably has other Webster (1994b)
affinities.
branches 5-angul h
-li-i u
i- ifi liti
i- I . I » i
placed in a relatively distant tribe, Jain. plicae
hair domatia on lower leaf surface {Tan- it
tealis); tuft
Dmi
Juts was not selected as outgroup, because
apc\
/,.'.','/.. jjai d- al of p.-fi.ili -l.'iidciU -.lilktd '<
many
Webster homoplasious
correct, parallel
[Vernicia cordata (Thunb.) Airy Shaw]; clavately if is
Mcnts would between .,>•;••:
exist !)<••
-talkt d 1' //'.. './ :::,in:<in,! Loir • I -
1 i
and Men: and wo likcK
Some characters, including several used in Web- siibtiibe i: ina< tin s< ild
.
\m
(1994b) keys, were not included in the phy- coii:pJi« ale die cladoiinun. feslm- ^liavy'- li\
's
would many more genera be
pothesis require that
logciiciii analysis, partly
and beyond
included in the analysis, that the
s(e.g.. is
\H
ope Wrbst.iV
of this article. .ugh ~ -
s< . i
. i i 1 1
and pubes cation was not based on phy logeia analyses, die
inflorescence in
fruit
;
m
sequence shows an inorplnes
of !axa rea.se in a <
(
anther dehii ;nce; ovary tapering gradually (for a list of synapomorphies, see Webster, 1994a).
a;.,
abruptly itilo style (important within Verni- ht j.i. is al to look for an outgroup in
i- i.
i . il
i I i
Umprn
petal venatio die -jibes treated bi lore tin- \1< n iliilcae.
),
It
*
-|
g |
I
^Hi=1i1i1liiHi
driulron Ricinorarpeae. subtribe Berlvinae)
(tribe
.mil I'iikk-roton |lnbe antorieae: former and bai-
\
(
I
&
known Chak-
ter as Fahrenheitia (Balakrishnan
rabarh. lWAj| nutgroup candidates, because
;.«tr
they have been well described in the lilerature and
The matrix comprised several polymorphic taxa.
abundant
mat. rial luis a\.nla,'!le !•• h«-. k Ii.iiae-
i .
MaHiladc
pn>\id.- pat-imoi, ..[ium-
lln- — -
• - t
zation for the polytypic characters by choosing one
The
state. other state is considered to be an auta-
pomorphy and counted as a step. The ng
is
>i!
,
are suggested:
i.|i|imizaln(iis
The rlade- c analvsis was peri'urnicd with PA P
I
A
Branch and
3.1.1 (Swofford, 1993).
Character 4 (pubescence of abaxial leal surface):
arch was performed with furthest addition
(sub)glabrous (state for Garcia (subtribe
1)
and Mulpais invoked. Nalistieal signifi-
ni tax.
i.
and Neoboutonia Neobou-
Garciinae) (subtribe
cance of the results was evaluated with the aid of
toninae); densely hairy (state 2) for Crotono-
a Bootstrap Anal\ >is a\ In <^ up to the level
•< !i
I «
gyne (subtribe Crotonogyninae).
=
D
6 (retaining trees five steps longer than the
all
Character 7 (shape of leaf blade base): cuneate
no- and Permutation
rsimonious a Tail
trees),
[i
Crotonogyne Crotonogyn-
(state 2) for (subtribe
The
Probability (FTP) test (Siddall, 1993). Boot-
and Decay were
strap (200 replicates) analyses also
PAUP
performed with 3.1.1 (Swofford, 1993), with
TBR,
Mul-
settings Heuristic search, Furthest,
for:
PTP
was performed conjunc-
pars. Mall's in
Si, test
'2
Charac-lei (nmnb.i ot ,lame,.s| \U -'air
If.
I.
Hennig86
tion with (Farris, 1988), with settings:
Crotonogyne and Manniophyton (sub-
2) for
characters unordered, nib*; bb*. Character opti-
Crotonogyninae).
tribe
Character 18 (ariaiigemenl of stamens): ordered in
A Maddison,
(Mii dis 1992).
i
two series (state 2) for Paracroton,
Several of the optimizations (Fig. can be in-
1)
The
terpreted differently. characters at the root are
parsiinorm cladograms were found now mli retell iilliti Ivpie.J for liie oiilgioiip
isl i|. a--
and are summarized in Borneodendron 13, 17, 23) or for the ingroup
(2,
=
(Fig. length 66, Consistency Index (CI) 0.67, see below. Characters 4 and 6 are regarded
1): (3, 12);
= =
RC
Retention Index (RI) 0.67, 0.45. All clad- as svnapomorpliies l-u -ml -.tribe Meuritinae with a
(ACCTRAN
mmiophv
ograms omprise suhtrihes
the lclie Aleii- reversal for Vernicia optimization);
i
P.tonogyninae, Garciinae, and Neobouton- racters can also be optimized as two par-
hi
ritil
inae. Subtribe Aleuritinae is sister group to the allel developments for Aleurites and Reutealis
DELTRAN
(DELTRAN
outgroup Paracroton, and these together form a optimization). optimization
.
im
(lad. w.lli pail "I -ill-tub.- ,ie--i ;th« iia-no- is used I'm character "2. a parallel level, .pin. Ioi
I til
_
: !> 'itiiliti, and Tapo- Hot », -.>,/< /,.•.';."' and /'./•.,- .',' 'lie alternative is a
-i i,
. i
;
ipliy for the ingroup with a reversal for
!n«
)i i|
,
ihrei former subtribes A leuritinae. uossennae. Paracroton). The state of character 19 is unknown
<
j
and mna.
Crotonogyninae), suhtril .m-ii -i-'m ',;;/«.-. r ibh ai lonog\ >uc,
lo; •;•.;,
j.- i(-.i > ): ti
group lo all other sublribes in tribe Aleiintiileae. ci. lliis rhaiactei is still regarded as a s\ nap. «tu<ir-
&
The Probability Tail Permutation test (Faith phv lor snl. irolone-Miinae, thus an apiciilale
lid...
<
Cranston, 1991; Kallersjo 1992) indicated connective
et al., is pi<
thai the data nuliix nlan -umiIk ml iii-mnl The strict consensus cladogram (Fig. 1) is sup-
-•
. .
=
=
of cladistic information (P 0.001001; a* ported by a relatively high CI (0.67) and RI (0.67).
0.000376). ! support rather poor. Only two
itistical is
i
A
strict consensus tree of all cladograms up to clades obtained a somewhat higher Deca] Index
69 steps, Decay Index 4, only resulted in a poly- I-.,- Kesulls) l!< ilslr ps in lin- study were never
95%
limn groups. sinel consensus ihe 16 or higher; therefore, no clades appeared to
I'm .ill \ <>i 1 1-
trees of up to 68 steps, Decay Index 3, showed have statistically significant support (Felsenstein,
1985). Recently, Hillis and Bull (1993, confirmed
\lei
up 67 Decay Index by K -hmo.
of to steps, lit\ I'i't.l} K «lrin
i - 1 r 1 * 1 1 : 1 1 1 \
1
9#-#
B!0JB
jj
O
B!uojnoqoeN(8K>&5-|
°>
I
J
I
III!
'^jii
XX##
B|0!U4©A iij
jjj
l ^
J
p
(8
f l j
k-b-o-I
CK>X-#
UOJOJOBJBd
&*5^
onslrated that bootstrap values of 70vf or higher versal (1(), stamens (> to Id); however, possesses
it
correspond probahilitv of 059r or more two unique autapomorphh
to a that a s ies 5-angulai
clade is real. In this interpretation subtribe Aleu- and sti-jjna- bilid "t inuliilid be -M, n.up re
I < .
ritinae (bootstrap TO'd bas siguilicanl support (Fig. between Reutealis and Xeuiu
-I
) 1 1 1 < i i| ,.
A
and none
of the other clades do. bootstrap supported by two parallel developments (though
1),
because corresponds
limit of ( ).Wf is very high, unique within subtribe Aleuritinae): 7 (leaf base
ll
to three apomorphies for a clade without any ho- cordate) and 16 (stamens 6
to 12).
moplasy (Felsenstein, 1985). Morphological data Monophyly of subtribes Crotonogyninae and
matrices usually do not include many ebaraclers.
runae supported in terms of unique
is
> 1
1
U
and. therefore. high bootstrap s ipporf eamiol be >-\ ib homji no -i-mIk up
el. » 1 .ir h >>!- \al
i
peeled chides. The bootstrap method pnre-
lor
all is ues or decay hid Sublri ireiinae mono-
is
l
statistical; docs not interpret the phv logenelie
l\ it generic, and, therefore, no bootstrap values or de-
We
significance of the characters. think that a clade
ca\ indices are available. seemingly has two
(,<in ui
with unique character combined with bootstrap
a a
unique character states (14, staminate petals 6 to
609f phvlogenetiealb acceptable hoof-
ol is (a (AV'A 13; 16, 63 to 160 stamens); however, character 16
corresponds one non-homoplasious
strap level to
applies only one ihe Iwo species die genus,
lo ol
ii:
character per chide!. niquc characters arc consul-
mmibei
I leaving only the higher slamiuale sepals
oi
unique
as a character.
m
being apomorphies.
elassificatioiis. reliable
Subtribe Grosserinae not supported the
is at all;
may
appear curious that subtribe Neobouton-
It
American Anomaloca-
relationships of the genera,
has bootstrap even
iiiae a of jusl (\Y/< thouiili is —
. it and
l\\ sjdl <-p<.i.il\
>,'i,i!,i in lie
,',':,.
i >l
supported by three apomorph proba
es his is
due
(both genera, vera luded aulapo
tn si noti-in.
hanges Neobou-
in b' In (•[!. i-i..ii- in
i l( ii i i .ii morphies, received the same coding). Within sub-
tonia (characters 3, hairs stellate; basal blade
...
tribe (hnsseririae. ( and Crusscm. bod'
'.tirticua \j
irdate leai bast- and 11, palmate ve-
li
ricau. eoustilule distim group (slippered !•
lln
'
i
Ml apomorphies
natiou). three 10. abaxial glan-
I and
urn eharact. 'K dots leaves; the
u< p< Ilia in
i
dular leaf hairs; ( onneeiive with gland; and 20,
>.
1 <
iion-ui iqin 17. -tameiis coi natej -aq.po^ d
ill '| |,,
anthers elongate) are unique: therefore, this lade
<
he|w..n
-hip .hi. />.••.•
In •;!,. a.
i« I. I. ..!.., i.i i
,
considered be phylogenetieally acceptable.
is to
M
hiiuun aid lagascar, Comores,
/ ,;.e; \1
.*. n. i.
The group Domohinea, Tannodia, and Tap i ,
<
weak
Borneo) (see ussion above)
is disc
has no significant phylogenelie vahie. be-
\~>V/<)
The
relationships between distinct groups
all
cause chaueler 12 incus)
noi imiijiie
1«> si:. is
!<> 1st
branches supported
(lower in Fig. are not b\
1)
within tribe Aleuritideae.
Decay
bootstrap values or Indices. Characters
1
Our purpose was to test the monophyly of sub-
and 22
(uionoec ions (danis) sepals) are
(5 pislillalt
determine which
tribe Yleutiliuae anil to generi.
unique: ul in
no*! iln\ 'c\ --i-e in -,.-\i Ia\a. .h.eai -
elassifieatioi! of subtribe Meuri'.uiac supported (
is
14 5 slamiuale and 22
We (3 to petals) (5 pistillate
chidistically. conclude that subtribe Aleuritinae
sepals), unique in Jus group, aie not uidqui n ihe
mi h\h Ji non Ji
is nof tn tb< h< itstr q> i- hi
lli<
.
Kuphorbiaeeae, nor are the inner eonnali sian
n>.
clade has a Decay Index of 3. and possesses a
it
divided pistillate disc as a unique char tc Mate
a.
One
of the outgroups, Paracroton, could not be
(character Airy Shaw's (1967) division of sub-
23).
placed parsimoniously die base ihe laiiogtam.
-e| ai
tllbi \ IflJt ll IU.1> 111') dlle,- i | t , I- -I pp..||- .|. . ' .,i <
an- -han-
least. Uciinti ai d \crni aeh have two unique b> i r \.ui..i- ,i| i pines with sub-
\ i • <
be Aleuritinae This suggests lhal tribe Xleurili-
lri
deae inonophvb'i
fruit: Vcrniruc H. glands in leaf margin is not ind lhal /'!•>,, '•>',"> is
The ma
m
ons; 25, seeds dorsally warty). Reutealis part of the ingroup. syna
is pi
>r
i
gh
untvpicalK characterized by single
a re-
Euphorbiaceae 66
tribe Aler steps, inch.
r ,gtl.
Decay
Indices higher
tl
Garden
Missouri Botanical
Candolle. Prodromus Svstematis Natural regni Yege-
is
lahihs 722 72
I5(2l: ,etori> Mass.,,, I'ans.
\ ,1 lilii.
1.
Paracroton).
K M On
Nixon. K. C. C: I'»>3. outgroups. Cla-
-titer.
J.
\imther s\napom<.rph> Meunlide.i.
tor
triht- lin-
9 413-120.
dislies
:
i\n,tri,>hm) character A &
i'llhliiiL. Is (slellate in. In Pax, F. K. Hoffmann PH2. In: \. Knrln (editor). Das
ineiituin). llus haia.l.r reverses several limes Pflanzenreieh IV.147.vi: 105-108. 110 121. 21.
nt figs.
I.
«
Engelmann,
cladogram and 2,\ 2.V Willi. Leipzig.
in the dot's not provide strong sup- I,,,
&
1914. In A. Engler (editor). Das Pflan-
port. Character states 2 (red exudate), 13 (staminate .
zenreieh lV.147.vii: 71-76, 10. Wilhelm Engel-
fig.
sepals absent), 17 (stamens connate), and 23
all
mann,
Leipzig.
ipishllate ahst
ills, ji. -,i«lrn apuiiii)iplii« s uphorbiaceae
I) In
(I
i i < i I
for the outgroup, Borneodendron, hecause these
-milk Mndagascan Euphorbiaceae
'::",.
\
states are exceplh oppo-iles. present I i i I I.
n. in. ll»
.1
I i On
the identity of I'aiu^-tuuwin and on alli.iihcs
tli«>
common.
in subtribe Aleuritideae, are However,
more taxa have he added the analysis 2'
to to a
l..r Hull. 0.32 ..12. I,,s 'tables '„,.,,'.
I.' !. 'l. 5. I.
heller evaluation inn/at ions, and
nl th.-s.- <-p|
lli.ii is
ideae-Aleurilideae-Crosserinae)
beyond scope with especial refer-
the of this study.
enee Madagascar, ami -uhsumplion
lo the ol Dnmt<lim,;i
Tins eladislic analysis, based morphul..-
<>n
! kew
Leandri. Bull. 53: 173-180.
characters, does not reveal whether or not tribe
Suldall. M. E. 1993. IVrmulation Tail Probability Test.
mnnophv m
Aleuritideae Hie addition
is lelii nl ire University of Toronto, Toronto.
.
taxa is warranted (or that goal.
I'osa. !«»«*•> ;,,...,„
It, ||, „
, , , 1
and Yrnnca (Euphorhiaeeae). Blumea 73-98.
44:
PUP:
IW3.
Swofford. Phvlogenetic \nalvsis sing
I). I.. I
Parsimony, Version 3.1.1. Illinois Natural History Sur-
i.wii.
(;,-,„.,
.
& Eupho
of the
the affinity of 1/ J. K. (,. For 11 ci a
I 1
393-395.
Shaw
'71. TapouU-s vHlamilii (Merr.) Airy
3032.
Icon. 37:
PI. lab.
&
The
N. Chakrabarty. 1995. genu,
n. P. T.
715-726.
48:
.
Notizbl. Bot. Gait. Berlin-
.
311-315.
I.
P.Taylor. 1950.
(
HIM
Sexualit
Monnecy
is often dillicull to diagnose because the
functionally unisexual inflorescences. For
this
rt
Stuppy
et al. (in press) recorded all three species
ii
Stony Brook. Nev nicia to be occasionally dioecious. However, this
in
ysis inonoecy was regarded as being typical for thest
cies. and the occasional unisexual specimen was
igi
His.' I).' M. « Bull. 1993. An empirical of young parts. Several
j. lest
i. , I
lootstrapping as a method for assessing confidence ryninae all ha\e lepi
in
.hvlogenelie ,,i,;iKs lv s vM |{„,|. 2: 182-192. the only liana in tribe Aleu-
|
.
&
Uannioplnl somewhat
llersjo, M.. S. Karris. A. Kluge C. Bull. 1992. rilideae. different: lepidole
J. (,.
Newness
and permutation. Cladislics 275-287.
8:
Kew
\n.erica: XI. Bull. Misc. Inform. 1933: 181 187.
2.
ig. 1.
indri. 1940. Sur genre malgaehe nouveau they a
J. 1111
fEuphorhineees. Bull. Sue. Bot. fiance 87: 270-285.
L
udell, C. 1945. The genus Garcia Vahl, a potential main pa p and, therefore, the
g - rl tl) , lil
I
source of superior hard quick-drying oil. Wrighiia I: air- ha\e been coded as l<.pidole. III. Iifle-Smith 198}?)
I.
I (
worded
we
lepidote hairs lUnoisiia: however, have
lot
en
onK
simple
hairs.
AnaKsis of Phylogeny and Character Ev Pubescence ol nbnxial leal -.urlacc: (sub)glabrous (I)
.
Sinauer. Sunderland.
3. \ln>si. husells. densely hairy
r
(2).
Millie, \,g,.Mensi>. J. |8(,i,. Euphorbia. This character was added because may provide a syn-
it
'
)
Volume Number
86,
and 6
y for Aleuritcs Rent, -ah;. I'll <•!>; acler to sepals: the other outgroup. I'aracroton, has 5 or 6
usualK be establish, d i;iul n oded as |)ol\morphic.
i i > i I | il . I . < <
mb
3-5
Only
ing parts. a few g. i i> , i nil II « I I inn it p i I ibsent (1), (or 6) (2),
some
ibe Neoboutonina. ha\ e species Petals are absent in the staminate flowers of Rornco-
•).
with glabrous leaves n n (kircia (sublribe
I
many
these genera are <<.<!<. the anahsis. Garciinae) is the only genus with (6-13) petals.
5. Glands on the petio h-al !>b h.c silion: absent There is a slight overlap in <-harael.-r slates, because some
! ( I
or presenl genera that usually have 5 petals can sometimes exhibit
(2).
Quite a few genera. nil- n sublribe variation Irorn "> lo () pelals within a particular inflores-
:,<!,
I t. . . i
"
' . species.
"in
[in
•I' ' .i >.. i .1 . > 'i
' i 1 |
top of the petiole. Most laxa possess Fusion of the staminate petals is consider.-. tvpi.nl foi
l I
The
sessile glands, but a few species have stalked o
lies.
shape ones
of the latter
could not be used in the analysis because the
onlv
,>re.
oded
therefore, /-,'..•; with a
th. is
(1) or often/always lobe d (2). ..ill i.| (
Lobed may
leaves pres [lewites
Of
and Vernicia in subtribe Meuntinae. all the ol her gen-
160
Manniophyton (3).
era, only fs> hi i« loi.o -V ind
Ml l<
i i
leave The coding of the states is difficult due to some overlap
Neoboulonia (subtribe N«•oboiilonmae) have lobed
and 2 and the Crotonogyne and Man-
in stales fact thai
rounded
or to cuneate, a 1
m:>i>!i::<.n (sublribe Crotonogyninae) are dimorphic for
or acute
(2).
J,,|,
v
st ||. | ,||„| III I . Ill III.K I lul s| It. s
unambiguous w
tribe Aleuritinae. State also ilhin the
1 is
other genera of tribe Ale uritideae, but state 2 variable
is
\ahl has (,.',- |(,() slam. -ns. (,. parnjlora Fundell ca. 2<>
and shows overlap anion; the species.
I
8. Marginal leaf glands: absent or abaxiall on the
(1), v
teeth (2), at teeth apices (3), or in teeth sinuse:> or ere-
, <Mb)
portant key character in W.-bsters I classification.
(
~"~"
ii
, ., .. 17- Fusion of s: in. ii . all !i- .- !. onl\ the inner whorl
Mosl laxa ol liiix \l. null. .lie n ind n !
. i. > I.
I
/W-
'
(
.leaf margins.
-
.1
i ,1 imlinae) he onlv taxon lha
, , , \\ v ,,. is
, ,
,
have glands on the abax.al s,de
of .1,
m
)
U|
^ ||r
, _ ,
homologous.
Vernicia
e
Meiirilinae) he onlv genus of tribe Aleuriti-
is
I
vhllJ, e
K!
-
I
leaves: ahsenl
I'.-lliic <i..|s in
.1 other two genera. Aleuritcs and
;
1
I'll,
i
found only
seri( . s (2 or or,|,. rt. in t or) 4
), ( | ;
(
mm
All members of subtribe Aleuritinae have palmate v.- ms „„.
wi ,|, ,. ,, , . ,,,.„ ,. , im ,. | 1;|U , ,| 1( . sl;im( .„ s
iMlmn. as.lo \hunu.n, ' mth-s
u o.d. n.l 2 I'aracroton the only
.,,,| in <u is
1I ,|
, ( , I
presence of leal lobes (character 0); leaves of Rcutealis or prolonged and apiculate
(3).
are never lobed. but have palmate v.-nalion. Triplinerved These three stales ma\ not he homologous. Prolongation
leaves, a special case ol pinnate venation, are found onlv „(' the connective (apiculate: present in subtribe Crolono-
Domolunai
in HI..V. ...- o| sublribe u\ M M „ia> be different from the presence of a gland
;„.)
i
Grossennae. (sublribe Neobouloninae). However, dividing these slates
12. Fusion of stanuii n nl oi fused, over one or two characters does not influence the results
il
i | I . i 1 I
rupturing spathaeeously
of the analysis.
(2).
mi
anthesis are tvpi eae gate, vermiform
' (2).
i I
The staminate sepals are free in I'araeroton anil ahsenl in Anthers with a short shape (1.5-2 times as long as
13. [Number of slamin 1 ill) 1(2). ml mi onlv iiisuhlrih. Voboiiton
it. <>i ,,, ,,:li
I [ i | i i I
6
or inae.
(3).
m
Sepals are absent one of the oulgioups. Borneo- 21. Tubercles of pollen exine: blunt oi sharp (2).
i.e..
( I )
derulron. All other genera of tribe Aleiu itideae show 2 lo Sharp exine tubercles are found onlv in Tannodia (in-
sepals x.ept la' uhliil b h <> nl nl i. .iinae) Hie tubercles
,<> .i
I . ( ' !
Garden
Missouri Botanical
SAMOA. Yumh;
(RISH). Ofu: G. 07, la (lilSH.
T.
SINGAPORE.
Dumat
N\). Pulau Sinclair
l-uut.
J.
5201 SO( ISLANDS.
IS).
[-,
II
I
I
SOLOMON
61154
Fosberg (HISH).
ahi, R.
F.
I
New
LANDS. Georgia: Tetrmara. BSIF (Maenu
TAIWAN.
Taitung: Tamali-lwia.m. lama
n _ ,n \u\ sliaw
ii li.
i (
MALAYSIA/SARAH. Lahad
ns):
2/605
(L);
am. St A ML 37836
Xleijer) (El. > \ far, tap
i i
*
Tembawan
Khl'
(L); Island. Wyatt-Srnilh)
(J
~
80510 SAX
27558
(L). Burgess) ('
(l>. F.
AUSTRAL 61836
ISLANDS.
Raivavae:
Raiurua,
&
John Fosberg 15683 (RISH. AUS- (L).
//. St. F. R. L).
TRALIA. Queensland: Unkelly, H viand 5406
B.
GO
(/Mid)
RRUNKI. BR( \ 541V Ka-ai: \lu<L, IVml.. ka'henga.
Herakas.
(L). I'ukol) ft.
111. (I.).
CAROLINE Deehamps 208 Orientals:
ISLANDS. Ponape: Isangi Yan-
M. (L). Terril..
Tolomail. Ta-
/
kamalsu 0H4 CHINA. Guangdong: gambi, Louis 2559 41 77 9676
(RISH). Guat.g- (L). (L). (L).
CAMEROON.
tonogyne
argentea
". 279 Guangxi: Rax: Ripinde: G.
G. Yip l.ungeliow.
B.
(I.), //.
.MW
/enke, 2029 2350 ,W7«
ILu
,-t,ln Si ' 7059,-! \\.\. ||,|. (I.I. .'*66tt ,/j.
il
•
GARON.
Libreville,
*:
/"
, ii
CONGO
Na.iua I. mi: Thakaundnm gii.i-gii De W Id.: (ZAIRE).
III
I
&
Degener F. Ordonez 14502 (NY). Viti Levu:
/W/ be
3600 Eeopoldville: Inongo l/opold
sinu. Gillespie (RISH). 1.). Territ.,
/. It. 1
FRENCH /4063
POLYNESIA.
C. tt/tert
II, (L).
ONCO
parOUoia
EMI) Croti.uogyiie |/\IRE|. Ogo-
H
I'ra.ri:
Fetionville.
VI 11. I
.
\H wib
Kaimann 6485
iiokm: ane.
(L).
isi •-,,.
i
CONCO
Kee l<>772 (RISH) INI>n\E- ggei Pax: (/AIRE). Orientu-
Yangambi, / 68/5
//wis
rr.. (L).
Ciolono^m
pr.u^ii liOON.
20800 Pax: Ripinde.
nas. hoarders \ \1
.S. //. ( 1 r,'.
Zenker 4666
mr: Ndeki. Kostermans (L).
1.
CAMKROON.
tkeri Pax: Lolodorf..lime
1913. /?ouvw *.n. (I): Ripinde. /.viAvy «22
(,'. (I.).
GABON.
Wiram)
Libreville. IR Dee. 1900, «. Klaine
IE P. s.n.
(/•: /
X- S.
.'
,
MADAGASCAR.
An
Simiulera errieri Leandri:
lara: liikluk.
I
I ! ./.
.
JAPAN.
Kvukvii: Okinawa. C. <r\llei:r,le6479{L).
(I.).
/•:. //.
.
FRENCH
GUYANA.
Walker 7750 LAOS. Dok Nanh. 1190 Rohr: Cayenne
(PS). Spire
.
MALAYSIA. 6459 MARTINIQUE.
(RM). Selangor: Ratu Caves, K. Ogata (L). St. Pierre.
•
I
SAN
10241a Sabah: Kulam
(L). Tennin, Ayer,
CONCO
maerantha
lArumpad) 15316 Sarawak: (ZAIRE)
(L). 1st Division, Telok «.r.,s,.-ra Pax: .-..p.dd-
I
MANGAREVA.
Pinang, S (B. Ue) 41942 (L). Duff. ville: M(i\en Kuiln Ten K.nvaka. Kevvango. R. De-
..
MARIANA
Fosberg II360(\\\S\\). ISLANDS. vred 2769
R.
F. (L).
Guam: 35533 MAR- Grossera major CAMEROON.
Yigo, R. Fosberg (RISH). Fax: June
F.
I...l..<l..rl.
QUESAS.
Uahuka. Quayle 1816
F. H. (BISH). 1919, C. d'Alleizelte s.n. R.p.mle. G. /.enke,
(I
):
MYANMAR
RM
F
(Rt Dukason 9394 1805 4227
A). H,.h„. G.
(L), (L).
CONGO
mullim.Os
Cross,-,;, Leonanl: |Z\IRE|.
\ «-,
( I.
II
3M4
Brass (NY). PHILIPPINES. Luzon: Uguna. Kivu: Kalehe km Kavumu-Wal-
Terr., vers 10 route
1
I,,s Banns. 1. I). F. F.lmer 17679 (L, MO, NY). ikale, Irangi, reserve IRSAC. G. Traupin 4529 (L).
PHOENIX
ISLANDS. 24627
Canton. Degener 4700
0.
(L).
PITCAIRN. CW1EROON.
Parker Valley. John 14980 Cross.ra pa.iic.dala
(I.). //. St. I'.iv Ripinde. G.
'
Volume Number
86,
kn 1711 ii /JYirXD.iwruil.i. :!{>.->0[\.).:u>y,n (N Lagiina. IIS Hi. C. McCrcgo, 22912 (B\1. k>:
.).
. ): i
3125 :«7^ ,'«77 :«60 FB (Aherns 2440 Minda-
(I.), .£?.?/ (I.). (I.). (L). Rizal, collector) (K. L).
P\H
3532 4920
(L). (L). nao: Agusan. Butaan. R. Metuloza 42223
tl).
)
CAMEROON
(\U
Arg.: LO (L); Davao. Santa Cruz. R. S. Williams 2852
km
Vleinl.a N
of l.-i.iHo.lisi.-ordilolia B...H <)\l.»i:i;s. M.iK.iudra?. C.
k
i (
6484
dWlleizette
(L).
Yaunde. G. Zenker 1404 (L); Bipinde, G. Zen&er I«- Cllamilii \in Shaw: MALAYSIA. Jo-
I\l
.): ii
I • i
CONGO
69fl (L). (ZAIKE). Orientale: Isangi IV.- hore: Mersing. Tenggaroh Eoresl Reserve, Compt.
KFP
Ymgnmbi.
Plateau Isaleui. 101. Ogata) 105167 Sabah: Sepilok
I.. /.o /,.v /4:>5.'i (I.). IK. (L).
./. (
\BO.N. Nvanga. C. dWlleizette 6460 Forest Beserve. SAN Wood) 16532
(I.). IG. 11. S. (L). iW.
CAMEROON. &
Aban) 128772 27676
Bip- Meijer IW. Meijer)
(L). (L):
I,-. <;. Zenker 1527 (I.). 3202a (I.). -fcM6 (L): \aun- Jalan Trig, Leila Forest Reserve, SAW (P. Lasan)
GABON.
//.W 62527
C. Zr/.Aer Libreville. C.
(I.). (L).
-\llri:,-i
hi In,
^
~p «im.
JAPAN.
(MEL). Hondo, 7138
E. H. Wilson (US):
Kyoto. Tagawa-Motozi 2091 kyushu, Isso-Rin-
(C);
do. K. Deguchi 5014 (\Y): No.no Saki. Piemt 925
VIETNAM.
Tonkin:
Hanoi. dWlleizette
(L). C.
6458a
(L).
W.-nieia
lordii ^ nlali\e
ll in ' .:•; i,ej.ie-i -.per
i i
CHINA.
Fujian: Yenping. Buongkang,
iniens): H. H.
Clump 3344 Geizhou:
(BO). Ts.invi Hsien. Liang
at: Cuming I'alung Nature Resent . on Air Pulih Feng Yah, A. N. Steward et al. 256 (NY). Guang-
dong: Lung Tou Shan. Ko 50228 NY). Gn-
S. P. (k.
&
.U//r/ie// /T«7« Kalimantan Selatan: Pleihari. angxi: Yung Hsien. Ta Te Tauen, A. N. Steward
(l.l.
Asam-Asam. bb 9494 Kalimantan Tengah: Cheo 1059 (NY). Hubei: Patung Hsien.
(L). K. C. C.
//.
NW
headwaters of Sungai Kehavan. 5 km of Turn- Chow 147 (NY). Hunan: Ping Ton Shan. IL T
bang Sian Logging Camp. S. Burle Tsang 23509 (US). Jiangsu: Nanking. Drum Tower.
J.
1121) ill. Kalimantan Timur: E. Ki C. Chiao 11993 (BO). Jiangxi: Lingehuan.
Y.
Y.
rajaan. \ of Sangkulirang. A. Koster il.i Tsiang 9822 (NY). Sichuan: Yulei. D. E. Bouffbrd
-
Snniatera Selatang: Bangka. I.obok-besar. Cuming 24035 (NY). Yunnan: Likiang. / Rock
et al. F.
& MM.USI
Pading. kmtermans Anta 927 5066 (NY. IS). Zhejiang: llangehow. Y Chiao
C.
\
1. (1 .1.
Pahang: S of Kuala Lompat. Krau (..line Besene.
kill' IHI iT. C. Whitman- 3580 22834 (kl. Utah Pradesh: Dehra Dun, U. Singh
) (\).
A\ INDONESIA. Jawa
Lokan Forest Reserve. Kinabalangan. S 131 (NY). Barat: Bangkelan.
Dist..
(J.
./.
UluLuak.Sun- G. B. Beumte A 913 (BO). JAPAN. Ryukvu: Oki-
gai Setap, S (0. Huron) 21312 (L). Pllll.il'PIM'.s nawa, Yahu, E. H. Walker el al. 6356 (L. US).
Ramos SOUTH kOHEA.
3791
IIS ( \. Yonglung. Ii. R. Yinger et al.
i
TMI
23828 Mindanao: Agusan Pro\ Tungao. TAIWAN. Chiavi Hsien: Fanlu Hsiang. Pan-
(Ll. (k).
..
R. Mendoza) 41965 (L). Palaivan: Pagdanan lienven. R. Lin 515 (MO).
II). Y.
SHI
(C. E. Rids-
,
THAILAM).
1511
Peninsular. Phallhal-
d(ile) (L).
Maxwell
Nature Park.
g F.
ill J.
I I t I-
86-547
(L).
Paracolon zevluiiicus (Vliill. Arg.) fialakr. & T. Chak- N. Hong Kon»: laipo. \
I
.
rah. (representative specimens): INDIA. Kerala: iMMi^l W.l
III II /, ' 1 I', ll
,
Cardamon Munnar-Alwaye
Road,
Hills. Pulivar. ea.
15 km from Munnar. C. E. Ridsdale 767 (L). SRI NESI\. Jawa Ti.iigah: Penwilemia,.. V
l,\Nk\. Kostermans Java An,
Call. Disi,-.. Iliuidiiiiiiia. 4. I2'>5 Tin.,..: Uj.-ml..,.
ll /,'.
I.
WaSlales:
(Bl i:\l\i 1',,,.,
I
Khant 1532't (Ki IPPINI -
/',. 1*1111 l.n/.
spe.-i.nens): INDONESIA. Jawa Barat: Tj.omas. C. na. LosBanos. Ill \. Catalan 20923 (B(
1 )
Hacker 36081 PHILIPPINES. Guimaras: Ho- LAND. Northern: Chiang Mai. Doi Su
A. (L).
(AM
M.agao. Luzon: Batangas. M. Ramos 1923 Maxicell 88 350 IE IN A
llo. (I.. L). \ VI.
.
MO): Cavite. Maragondong, BS Fenix) 5160 Wun; \|„ Mian. Tana 28987
IF. la.ii
II T. l\