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Recommended citation: APG III (2009). This paper was compiled by Birgitta Bremer, Kre Bremer, Mark W.
Chase, Michael F. Fay, James L. Reveal, Douglas E. Soltis, Pamela S. Soltis and Peter F. Stevens, who were
equally responsible and listed here in alphabetical order only, with contributions from Arne A. Anderberg, Michael
J. Moore, Richard G. Olmstead, Paula J. Rudall, Kenneth J.
Sytsma, David C. Tank, Kenneth Wurdack, Jenny Q.-Y. Xiang and Sue Zmarzty (in alphabetical order). Addresses: B.
Bremer, The Bergius Foundation at the Royal Swedish Academy of Sciences, PO Box 50017, SE-104 05 Stockholm,
Sweden; K. Bremer, Vice Chancellor, Stockholm University, SE-106 91 Stockholm, Sweden; M. W. Chase, M. F. Fay,
Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3DS, UK; J. L. Reveal, L.H. Bailey
Hortorium, Department of Plant Biology, 412 Mann Building, Cornell University, Ithaca, NY 14853-4301, USA; D.
E. Soltis, Department of Biology, University of Florida, Gainesville, Florida 326118525, USA; P. S. Soltis, Florida
Museum of Natural History, University of Florida, Gainesville, Florida, 326117800, USA; and P. F. Stevens,
Department of Biology, University of Missouri-St. Louis and Missouri Botanical Garden, PO Box 299, St. Louis,
Missouri 631660299, USA
Received 12 August 2009; accepted for publication 18 August 2009
A revised and updated classification for the families of flowering plants is provided. Many recent studies have yielded increasingly
detailed evidence for the positions of formerly unplaced families, resulting in a number of newly adopted orders, including
Amborellales, Berberidopsidales, Bruniales, Buxales, Chloranthales, Escalloniales, Huerteales, Nymphaeales, Paracryphiales,
Petrosaviales, Picramniales, Trochodendrales, Vitales and Zygophylla-les. A number of previously unplaced genera and families are
included here in orders, greatly reducing the number of unplaced taxa; these include Hydatellaceae (Nymphaeales), Haptanthaceae
(Buxales), Peridiscaceae (Saxifragales), Huaceae (Oxalidales), Centroplacaceae and Rafflesiaceae (both Malpighiales), Aphloiaceae,
Geisso-lomataceae and Strasburgeriaceae (all Crossosomatales), Picramniaceae (Picramniales), Dipentodontaceae and
Gerrardinaceae (both Huerteales), Cytinaceae (Malvales), Balanophoraceae (Santalales), Mitrastemonaceae (Ericales) and
Boraginaceae (now at least known to be a member of lamiid clade). Newly segregated families for genera previously understood to
be in other APG-recognized families include Petermanniaceae (Liliales), Calophyllaceae (Malpighiales), Capparaceae and
Cleomaceae (both Brassicales), Schoepfiaceae (Santalales), Anacampserotaceae, Limeaceae, Lophiocarpaceae, Montiaceae and
Talinaceae (all Caryophyllales) and Linder-niaceae and Thomandersiaceae (both Lamiales). Use of bracketed families is abandoned
because of its unpopu-larity, and in most cases the broader circumscriptions are retained; these include Amaryllidaceae,
Asparagaceace and Xanthorrheaceae (all Asparagales), Passifloraceae (Malpighiales), Primulaceae (Ericales) and several other
smaller families. Separate papers in this same volume deal with a new linear order for APG, subfamilial names that can be used for
more accurate communication in Amaryllidaceae s.l., Asparagaceace s.l. and Xanthorrheaceae s.l. (all Asparagales) and a formal
supraordinal classification for the flowering plants. 2009 The Linnean Society of London, Botanical Journal of the Linnean
Society, 2009, 161, 105121.
105
106
Plant
Book,
Mabberley
(2008: xi, 927,
929) consulted
widely among
tax-onomists
about
which
alternative they
preferred, and,
more recently,
the issue of
these
alternative
circumscription
s was discussed
by researchers
representing
several
European
herbaria (e.g.
2.
K, E, BM, P, G
and the Dutch
herbaria
collectively)
that are in the
process
of
reorganizing
their collections
3.
along
APG
lines.
They
have
all
agreed
to
adopt APG III
as
their
standard and
the
linear
order
of
Haston et al.
(2009).
In
general,
the
broader
circumscriptions
have
been
favoured and
are
adopted
here.
Papers
over
the last few
years
have
clarified the
positions
of
isolated
families
including
Ceratophyllaceae,
Chloranthacea
e
and
Picramniaceae
(Jansen et al.,
2007; Moore
et al., 2007;
Wang et al.,
2009), and this
has
necessitated
addition
of
orders
not
previously
recognized by
APG.
A
few
genera/familie
s, members of
which
had
either not been
sequenced
before or for
which
chimaeric
sequences
were
produced,
were wrongly
placed. Thus,
families like
Guamatelacea
e have been
added;
Guamatela
used to be
placed
in
Rosaceae, but
molecular data
places it in
Crossosomatal
es (Oh &
Potter, 2006).
Hydatel-laceae
have
been
moved from
Poales
to
Nymphae-ales
(Saarela et al.,
2007).
4.
There are a
few
cases
where
the
general pattern
of
relationships
has
not
changed much
since APG II,
but
our
appreciation of
the pattern of
variation has.
For example,
this
helps
justify
inclusion
of
Ixerbaceae in
Strasburgeriac
eae.
5.
Finally, a few
family
circumscriptio
ns suggested
by APG II did
not
reflect
general usage,
so we have
modified
these,
an
example being
the
broadly
circumscribed
Brassicaceae,
which are here
split into three
families.
In general, we
have tried not to
phylo-genetic
uncertainty that
may necessitate
further revision of
familial or ordinal
limits. For further
dis-cussion on the
variation in the
groups discussed,
potential
apomorphies, etc.,
see the literature
cited and Stevens
(2001);
particularly
important recent
work
includes
Wang et al. (2009:
rosids), Tank &
Donoghue
(in
press), Moore et
al. (2008, in press:
core
eudicots),
Wurdack & Davis
(2009:
Malpighiales) and
Linnean
Botanical Journal of the
Society of Linnean Society, 2009,
London,
161, 105121
ton
et al.
h (200
e 9).
g With
e in
n order
er s, the
al sequ
se ence
q of
u famil
e ies is
n alpha
c betic
e al.
ofSym
or bols:
d *new
er famil
s y
fo place
ll o ment
w;
s new
th ly
e reco
le gnize
ft d
to order
ri for
g the
ht APG
se syste
q m;
u new
e famil
ny
c circu
e mscr
ofiptio
th n
e descr
la ibed
rg in
el the
y text;
la $fam
d ilies
d that
er repre
iz sent
e the
d broa
tr der
e circu
e mscr
in iptio
H n of
as optio
ns
avail
able
in
APG
II
and
favo
ured
here,
$
$fam
ilies
that
were
in
squar
e
brac
kets
in
APG
II,
the
narro
wer
circu
mscr
iptio
ns
favo
ured
here.
The
list
refle
cts a
starti
ng
date
for
all
flow
ering
plant
famil
y
name
s as
4
Aug
ust
1789
(Juss
ieu,
Gene
ra
plant
arum
).
Full
ci esent
ta -ing
ti the
o relati
n onshi
s ps
ar amo
e ng
a the
v majo
ai r
la grou
bl ps
e reco
el gnize
se d
w here
h is
er prese
e nted
( in
R Figur
e e 1.
v
e
al
,
2 CL
0 AS
0 SIF
8 IC
ATI
o ON
n OF
w FL
ar O
d) WE
. RI
A NG
s PL
u AN
m TS
m
Am
ar
borel
iz
lales
e Meli
d kyan
p,
h A.V.
yl Bobr
o ov &
g Zayt
e zeva
n (199
et 9)
ic
tr A
e m
e bo
re rel
pr lac
ea
e
Pi
ch
on
(1
94
8),
no
m.
co
ns.
Th
e
evide
nce
that
Amb
orell
acea
e are
sister
to all
other
angi
osper
ms is
clear
(e.g.
Hans
en et
al.,
2007
;
Janse
n et
al.,
2007
;
Moo
re et
al.,
2007
).
How
ever,
even
if
they
were
sister
to
Nym
phae
ales
(e.g.
Gore
myki
n,
Viola
&
H xon
el with
l out
w chara
ig cters.
,
2
Ny
0
m
0
ph
9)
ae
,
ale
th
s
e
Sa
y
lis
s
b.
h
ex
o
Be
ul
rc
d
ht.
b
&
e
J.
k
Pr
e
esl
pt
se (1
p 82
ar 0)
at $
e $C
as ab
th o
ei m
ba
r
in ce
cl ae
u Ri
si ch
o .
n ex
in A.
N Ri
y ch
m .
p (1
h 82
a 2),
e no
al m.
es co
w ns.
o *
ul H
d yd
re ate
s lla
ul ce
t
ae
in U.
a Ha
ta m
an
n
(1
97
6)
$
$
N
y
m
ph
ae
ac
ea
e
Sa
lis
b.
(1
80
5),
no
m.
co
ns.
Th
ere
seem
s to
be a
gene
ral
prefe
rence
for
keepi
ng
Cabo
mbac
eae
and
Nym
phae
acea
e
separ
ate,
altho
ugh
both
are
small
in
term
s of
speci
es.
The
two
famil
ie e
s was
ar unex
e e pecte
as d,
il but it
y is
c well
h supp
ar orted
a morp
ct holo
er gical
iz ly
a and
bl embr
e. yolo
T gical
h ly
e (Saar
in ela
cl et
u al.,
si 2007
o;
n Fried
ofman,
H 2008
y;
d Ruda
at ll et
el al.,
- 2008
la ).
c
e
a
e
(p
re
vi
o
u
sl
y
in
cl
u
d
e
d
in
P
o
al
es
)
h
er
Aust
ro
ba
ile
ya
le
s
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kh
t.
ex
R
ev
ea
l
(1
99
2)
A
us
tr
ob
ail
ey
ac
ea
e
Cr
oi
za
t
(1
94
3),
no
m.
co
ns
.
$S
ch
is
an
dr
ac
ea
e
Bl
u
m
e
(1
83
0),
no
m.
co
ns
.
(i
nc
lu
d
-
7
)
,
n n
g o
I m
l .
l
i c
c o
i n
a s
c .
e
a S
e c
A h
. i
C s
. a
S n
m d
., r
n a
o c
m e
. a
c e
o
n s
s .
. l
) .
T
r a
i r
m e
e
n w
i e
a l
c l
e
a c
e h
L a
. r
S a
. c
G t
i e
b r
b i
s z
( e
1 d
9 .
1
Chl
orant
hales
R.Br.
(183
5)
C
hl
or
an
th
ac
ea
e
R.
Br
.
ex
Si
m
s
(1
82
0),
no
m.
co
ns
.
Ch
loran
thace
ae
are
prob
ably
sister
to
mag
nolii
ds
(Mo
ore
et
al.,
2007
).
Sepa
rate
ordin
al
statu
s is
warr
anted
beca
use
of
their
phyl
ogen
etic
posit
io l
n; l
th a
e c
y e
ar a
e e
al
s M
o a
mr
or t
p .
h
(
ol
1
o
8
gi
3
c
2
al
)
ly
,
di
st n
in o
ct m
iv .
e.
B
e
r
c
h
t
.
&
J
.
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r
e
s
l
(
1
8
2
0
)
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r
c
i
o
MAGNOLIIDS
s
n
t
Ca s
o
n.
l
e W
o
int
l
c
er
l
h
ac
a
i
ea
l
e
a
e
R.
c
s
Br
e
.
a
C ex
e
r
Li
o nd
J
n l.
u
q (1
s
u 83
0)
s
i
,
.
s
no
t
m.
(
co
1
(
ns
7
1 .
8
9
Pi
9
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)
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,
) e
r
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.
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n
g
l
.
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y1
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a ,
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gn
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r .
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hP
i
( p
1e
8r
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, a
e
n
oG
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. s
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. 1
7
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o
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.
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relati
onshi
ps of
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nora
ceae
are
uncle
ar
withi
n
Piper
ales.
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u
r
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u
s
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.
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r
.
(
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r n
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&a
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(
1
8
1
9
)
,
A
t n
ho
e m
r .
o
s c
po
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r s
m.
a
t G
a o
c m
e o
a r
e t
e
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. a
c
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e
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e
i
c
h
e
(
1
8
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6
)
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o
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.
c
o
n
s
.
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e
r
n
a
n
d
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a
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l
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m
e
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1
8
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6
)
,
n
o
m
. (
1
c 8
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s )
. ,
Ln
a o
um
r .
a
c c
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e s
.
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uS
s i
s p
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r
( u
1n
7a
8c
9e
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, e
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mh
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oe
n
s (
. 1
9
M7
o0
n)
i
Mag
m
no
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lia
a
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s
e
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u
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s
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er
ch
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&
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es
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(1
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nn
on
ac
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e
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ss.
(1
78
9),
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m.
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ns
.
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eg
en
eri
ac
ea
e
I.
W
.B
ail
ey
&
A.
C.
S
m.
(1
94
2),
no
m.
c
o
n
s
.
E
u
p
o
m
a
t
i
a
c
e
a
e
O
r
b
.
(
1
8
4
5
)
,
n
o
m
.
c
o
n
s
.
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i
m
a
n
t
a
n
d
r
a
c
e
a
e
D
i
e
l
s
(
1
9
1
7
)
,
n
o
m
.
c
o
n
s
.
M
a
g
n
o
l
i
a
c
e
a
e
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u
s
s
.
(
1
7
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9
)
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n
o
m
.
c
o
n
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.
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y
r
i
s
t
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.
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r
.
(
1
8
1
0
)
,
n
o
m
.
c
o
n
s
.
MONOCOTS
Acor
ales
Link
(183
5)
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ra
ce
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art
in
ov
(1
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m
at
al
es
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ex
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er
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Pr
es
l
(1
82
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is
m
at
ac
ea
e
V
e
nt
.
(
1
7
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9
),
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o
m
.
c
o
n
s.
(i
n
cl
u
di
n
g
i
m
n
o
c
h
a
r
i
t
a
c
e
a
e
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a
k
h
he
Li
n
2 ne
0 an
0S
9 oc
Tie
ty
t
.
)
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e
x
n
o
m
.
C
r
o
n
q
u
i
s
t
)
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p
o
n
o
g
e
t
o
n
a
c
e
a
e
P
l
a
n
c
h
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(
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8
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6
c
o
n
s
.
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r
a
c
e
a
e
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u
s
s
.
(
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7
8
9
)
,
n
o
m
.
c
o
n
s
.
of anical
L Journal
o of the
n Linnean
d Society,
o 2009,
n, 161,
B 105
ot 121
108
angiosperms
monocots
eudicots
asterids
Amborellales
Nymphaeales
Austrobaileyales
Piperales
Canellales
magnoliids
Magnoliales
Laurales
Chloranthales
Commelinales
Zingiberales
commelinids
Poales
Arecales
Dasypogonaceae
Asparagales
Liliales
Pandanales
Dioscoreales
Petrosaviales
Alismatales
Acorales
Ceratophyllales
Ranunculales
Sabiaceae
Proteales
Buxales
Trochodendrales
Gunnerales
Cucurbitales
Fagales
Rosales
Fabales
fabids
Celastrales
Oxalidales
Malpighiales
Zygophyllales
Malvales
Brassicales
Huerteales
Sapindales
malvids
Picramniales
Crossosomatales
Myrtales
Geraniales
Vitales
Saxifragales
Dilleniaceae
Berberidopsidales
Santalales
Caryophyllales
Cornales
Ericales
Garryales
Gentianales
lamiids
Lamiales
Solanales
Boraginaceae
Aquifoliales
Escalloniales
Asterales
campanulids
Dipsacales
Paracryphiales
Apiales
Bruniales
t
A ers andor
Bayesian
posterior
P some
probabilities greater than
Figure
1.
G families 0.95 in large-scale analyses
Interrelations
supported of angiosperms. See text for
hips of the
I by
literature supporting these
APG
III
I jackknife/ relationships.
Newlyorders
and
I bootstrap recognized-for-APG orders
some families
o percentag are denoted (). Some
supported by
r es greatereudicot families not yet
jackknife/boo
d than 50classified to order are not
shown.
Tn
of London, Botanical Society, 2009,
hSocietyJournal of the Linnean 161, 105121
S
c
h
e
u
c
h
z
e
r
i
a
c
e
a
e
h
t
.
(
1
9
9
5
)
Zo
ste
ra
ce
ae
D
u
m
ort
.
(1
82
9),
no
m.
co
ns.
F
.
R
u
d
o
l
p
h
C
i
onvi
(
ncin
1
g
8
evide
3
nce
0
for
)
the
,
mon
n
ophy
o
ly of
m
Alis
.
mata
c
ceae
o
s.s.
n
is
s
lacki
.
ng
T
(e.g.
o
fChen
iet
eal.,
l2004
da, b)
iand
athe
cfamil
ey
adoes
enot
have
Tany
aapo
kmorp
hies.
Whe
n
com
bine
d
with
Limn
ocha
ritac
eae,
a
famil
y
that
was
only
relati
vely
recen
tly
descr
ibed
(Cro
nquis
t,
1981
), the
enlar
ged
famil
y has
sever
al
disti
nctiv
e
chara
cters.
It
may
be
nece
ssary
to
split
off
Mau
ndia
from
Junc
agina
ceae
(Iles
et
al.,
2009
; S.
von
Meri
ng &
J. case
Wto
. creat
Ke a
a large
dr
er singl
ei e
t, famil
p y for
er the
s. large
c r
o clade
m.
m More
.), study
a is
n need
d ed
Mbefor
a e
u anot
n her
di mon
a ogen
c eric
e famil
a y is
e reco
N gnize
a d in
k Alis
ai mata
is les.
a
v
ai
la Petr
bl osavi
ales
e.
Takh
H t.
o (199
w 7)
e Pe
v tro
er sa
vi
,
it ac
m ea
e
ig
H
ht
ut
b ch
e .
b (1
et 93
te 4),
no
r
in m.
th co
ns.
is
Th
e
isolat
ed
posit
ion
of
Petro
savia
ceae
here
is
well
supp
orted
(e.g.
Tam
ura
et
al.,
2004
;
Chas
e et
al.,
2006
),
henc
e its
ordin
al
statu
s.
Dios
corea
les
R.Br.
(183
5)
B
u
r
m
a
n
n
i
a
c
e
a
e
B
l
u
m
e
i
( a
1 c
8 e
2 a
7 e
)
, F
n r
o .
m
. e
c x
o
n B
s j
. u
D r
i z
o o
s n
c
o (
r 1
e 8
a 4
c 6
e )
a
e Th
ismia
ceae
R
.J.Ag
ardh
B
(185
r
8)
.
may
(
turn
1
out
8
to be
1in a
0clade
)separ
,ate
nfrom
oBur
m
man
.niace
cae
oand,
nsimil
sarly,
.the
N
morp
aholo
rgical
tly
hdisti
enctiv
ce
Tacc
acea
e
Dum
ort.
(182
9),
nom.
cons.
,
from
Dios
corea
ceae.
Phyl
ogen
etic
relati
onshi
ps in
Dios
corea
les
that
supp
ort
such
chan
ges
have
been
foun
d by
Merc
kx et
al.
(200
6)
and
Merc
kx &
Bida
rtond
o
(200
8),
and
Merc
kx et
al.
(200
9)
even
sugg
est
that
This
miac
eae
s.s.
may
b we
e have
p been
ar cons
a ervat
p ive
h and
yl not
et adop
ic ted
. any
G alter
iv ed
e cirn cums
th cripti
e ons
pr at
o this
bl stage
e .
m
s
in Pan
u d
n a
d n
er a
st l
a e
n s
di R
n .
g B
re r.
la e
ti x
o B
n e
s r
hi c
p h
s t.
of &
m
y J
c .
o P
h r
et e
er s
ot l
ro (
p 1
hi 8
c 2
gr 0
o )
u C
p y
s, c
l
a
n
t
h
a
c
e
a
e
P
o
it
.
e
x
A
.
R
i
c
h
.
(
1
8
2
4
),
n
o
m
.
c
o
n
s.
P
a
n
d
a
n
a
c
e
a
e
R
.
B
r.
(
1
8
1
0
),
n
o
m
.
c
o
n
s
.
S
t
e
m
o
n
a
c
e
a
e
C
a
r
u
e
l
(
1
8
7
8
)
,
n
o
m
.
c
o
n
s
.
T
r
i
u
r
i
d
a
c
e
a
e
G
a
r
d
n
e
r
(
1
8
4
3
),
n
o
m
.
c
o
n
s
.
V
e
ll
o
z
i
a
c
e
a
e
J
.
A
g
a
r
d
h
(
1
8
5
8
),
n
o
m
.
c
o
n
s
.
Lilial
es
Perle
b
(182
6)
A
l
s
t
r
o
e
m
e
r
i
a
c
e
a
e
D
u
m
o
r
t
.
(
1
8
2
9
)
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n
o
m
.
c
o
n
s
.
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i
n
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l
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d
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g
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z
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r
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e
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o
t
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y
)
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a
m
p
y
n
e
m
a
t
a
c
e
a
e
D
u
m
o
r
t
.
(
1
8
2
9
)
C
o
l
c
h
i
c
a
c
e
a
e
D
C
.
(
1
8
0
4
)
,
n
o
m
.
c
o
n
s
.
C
o
r
s
i
a
c
e
a
e
B
e
c
c
.
(
1
8
7
8
)
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n
o
m
.
c
o
n
s
.
L
i
l
i
a
c
e
a
e
J
u
s
s
.
(
1
7
8
9
)
,
n
o
m
.
c
o
n
s
.
M
el
an
thi
ac
ea
e
B
at
sc
h
ex
B
or
kh
.
(1
79
7),
no
m.
co
ns
.
*P
et
er
m
an
ni
a s.
c Ri
e po
a go
e na
H ce
u ae
t C
c on
h ra
. n
( &
1 Cl
9 iff
3 or
4 d
) (1
, 98
n 5)
o S
m mi
. la
c ca
o ce
n ae
s Ve
. nt.
P (1
h 79
i 9)
l ,
e no
s m.
i co
a ns
c .
e
a
Pe
e
term
D
annia
u
ceae
m
are
o
morp
r
holo
t
gical
.
ly
(
and
1
phyl
8
o2genet
9icall
)y
,disti
nnct.
oLuzu
m
riaga
.ceae,
cconsi
osting
n
of
two
small
gene
ra
with
gene
raliz
ed
lily
floral
morp
holo
gy,
are
sister
to
Alstr
oeme
riace
ae
and
have
the
same
disti
nctiv
e
twist
ed
petio
les,
so
com
binat
ion is
in
order
(see
also
Mab
berle
y,
2008
).
Aspa
ragal
es
Link
(182
9)
$
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r
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D
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m
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h
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ns
.
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co
ph
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ce
ae
Le
yb
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(1
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ns
.
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an
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. or
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8 82
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M
ae
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Allia
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C
are
hreco
agnize
sd
ehere
,(Am
R
arylu
lidac
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eae
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l
recen
l
tly
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erve
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.d
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.Allia
Fceae,
adespi
yte
Allia
(ceae
2bein
0g the
0older
0name
)).
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ral
charn acter
e s
x supp
p ort
a the
n com
d bine
e d
d grou
A p.
m Agap
ar antha
yl ceae,
li if
d kept
a separ
c ate,
e are
a weak
ly
chara
cteri
zed;
the
famil
y is
mon
ogen
eric.
Th
e
area
arou
nd
Aspa
ragac
eae
is
diffic
ult
from
the
stand
point
of
circu
mscr
iptio
n.
Alth
ough
Aspa
ragac
eae
s.l.
are
heter
ogen
eous
and
poorl
y
chara
cteri
zed,
Aspa
ragac
eae
s.s.,
Agav
acea
e,
Lax
man
niace
ae,
Rus2 009 The
Loc
i ie
n ty
n of
eL
ao
nn
Sd
o Journal
n, of the
B Linnean
ot Society,
a 2009,
ni 161,
c 105
al 121
110
nom. cons.
Joinvilleace
ae Toml. &
A.C.Sm.
(1970)
Juncaceae
Juss. (1789),
nom. cons.
Mayacaceae
Kunth
(1842), nom.
cons.
R
e
s
t
i
o
n
a
c
e
a
e
Poa
cea
e
Bar
nha
rt
(18
95),
no
m.
con
s.
Rap
atea
cea
e
Du
mor
t.
(18
29),
no
m.
con
s.
R
.
B
r
.
)
,
n
o
m
.
c
o
n
s
.
Typhaceae
Juss. (1789),
nom. cons.
(including
Sparganiacea
e Hanin,
nom. cons.)
Xyridaceae
C.Agardh
(1823), nom.
cons.
(
1
8
1
0
)
,
Sparganiaceae
are included in
Typhaceae;
the
two families are
monogeneric,
occupy
similar
habitats and share
a
number
of
features. That they
were
treated
separately in APG
II was a mistake
(M. W. Chase,
pers.
comm.).
They have in the
past
been
combined;
Mabberley (2008)
suggested
that
their combination
would be in order.
n
o
m
.
c
o
n
s
.
T
h
u
r
n
i
a
c
e
a
e
E
n
g
l
.
(
1
9
0
7
Zi
n
g
i
b
e
r
a
l
e
s
G
r
i
s
e
b
.
(
1
8
5
4
)
C
a
n
n
a
c
e
a
e
J
u
s
s
.
(
1
7
8
9
)
,
n
o
m
.
c
o
n
s
.
C
o
s
t
a
c
e
a
e
N
a
k
a
i
(
1
9
4
1
)
H
e
l
i
c
o
n
i
a
c
e
a
e
V
i
n
e
s
(
1
8
9
5
)
L
o
w
i
a
c
e
a
e
R
i
d
l
.
(
1
9
2
4
)
,
n
o
m
.
c
o
n
s
.
Mara
ntace
ae
R.Br.
(1814
),
nom.
cons.
Musa
ceae
Juss.
(1789
),
nom.
cons.
Strelit
ziace
ae
Hutch
.
(1934
),
nom.
cons.
Zingi
berac
eae
Marti
nov
(1820
),
nom.
cons.
PROBAB
LE
SISTER
OF
EUDICO
TS
Ceratophyllales
Link (1829)
Ceratophyllace
ae Gray (1822),
nom. cons.
The molecular
evidence
that
Ceratophyllaceae
are
sister
to
eudicots
is
becoming clearer
(Jansen et al.,
2007; Moore et
al., 2007, but cf.
Goremykin et al.,
Ranunculales Juss.
ex Bercht. &
J.Presl (1820)
Berberidaceae
Juss. (1789),
nom. cons.
$Circaeasterace
ae Hutch.
(1926), nom.
cons.
(including
Kingdoniaceae
Airy Shaw)
Eupteleaceae
K.Wilh. (1910),
nom. cons.
Lardizabalacea
e R.Br. (1821),
nom. cons.
Menispermacea
e Juss. (1789),
nom. cons.
EU$Papaveraceae
Juss. (1789),
DIC
nom. cons.
OTS
(including
Linnean
2009 The Society of
London,
Fumariaceae
Marquis,
nom. cons.,
Pteridophyl-
laceae Nakai
ex Reveal &
Hoogland)
Ranunculaceae
Juss. (1789),
nom. cons.
We adopt broad
limits
for
Circaeasteraceae
and Papaveraceae,
as
this
is
commonly done
(Judd et al., 2007;
Mabberley, 2008),
and
the
two
families are well
characterized in
their
broader
circumscriptions.
The two families
into
which
Circaeasteraceae
have been divided
(Circaeasteraceae
and
Kingdoniaceae)
r
e
s
l
n
a
c
e
a
( e
1
8T
2.
0L
) e
s
Nt
ei
l b
u.
m
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n8
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i
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ho
. n
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( .
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8$
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ne
oa
mc
. e
a
ce
o
nJ
s u
. s
s
$.
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l 1
a7
t 8
a9
)
,
n
o
m
.
c
o
n
s
.
Pl
atana
ceae,
altho
ugh
mon
ogen
eric,
are
morp
hologic
ally
disti
nct
from
Prote
acea
e,
and
the
two
have
neve
r
been
com
bine
d
previ
ousl
y;
mem
bers
of
the
broa
der
famil
y
woul
d
have
few
featu
res
in
c
o
m
m
o
n.
m.
co
ns
.
(
i
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d
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g
Tr
o
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h
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ro
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ro
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h
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c
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s
e
.
a
)
e
.
E
ic
hl
A
er
sepa
(1
rate
8
6 orde
5)r for
, this
n mor
o phol
ogic
ally
disti
nct
clade
is
warr
ante
d;
the
two
mon
ospe
cific
gene
ra in
Troc
hode
ndra
ceae
s.l.,
Tetr
acen
tron
and
Troc
hodend
ron,
have
muc
h in
com
mon.
Bu
xales
Takh
t. ex
Reve
al
(199
6)
$
B
u
x
a
c
e
a
e
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u
m
o
r
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.
e
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8 el
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n
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) 00
, 2)
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o T
he
m
.limit
cs of
oBux
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se are
.expa
(nded
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nmon
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uDidy
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nhave
gthe
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inctiv
de
ypolle
m
n
eand
lche
amistr
cy as
eat
aleast
epart
of
LBux
eacea
ae,
naltho
dugh
rthere
iis
)curre
*ntly
H
no
aevid
p
ence
t
afor
nthe
tpara
hphyl
ay of
cthe
elatter
a.
Som
e
morp
holo
gical
featu
res
sugg
est
that
Hapt
anth
acea
e are
best
place
d
here,
but
they
are
disti
nct
from
all
other
angi
ospe
rms
(Dou
st &
Stev
ens,
2005
). An
order
for
the
two
famil
ies is
warr
ante
d.
Note
that
relati
onsh
ips
of
Troc
hode
ndral
es
and
Buxa
les
rema
in
uncl
ear,
al .
th
oe
ux
g
hR
the
ev
ye
ara
el
c
er(
ta
1
in
9
ly
9
to
2
b
)
e
pl
$
a
c$
eG
du
inn
thn
is e
pr
ara
t c
oe
f a
the
e
tr M
ee
e. i
s
n
.
$
$
M
y
r
o
t
h
a
m
n
a
c
e
a
e
N
i
e
d
.
(
1
8
9
1
)
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n
o
m
.
c
o
n
s
.
(
CORE
1
EUDICOTS
T
8
4
u2
n)
n,
e
r n
ao
l m
e.
s
c
To
an
ks
h.
t
he
two
famil
ies
share
no
impo
rtant
featu
res
and
so
are
kept
separ
ate,
al T
th his
o fami
u ly
g has
h no
b stabl
ot e
h posit
ar ion
e as
m yet
(Mo
o
ore
n
et
o
al.,
g
in
e
press
n
).
er
The
ic ordi
. nal
nam
D
ie,
lDille
lniale
e
s
n
iDC.
aex
cBerc
eht. &
aJ.Pre
esl, is
avail
S
aable.
Sal
ix
si
b
f
.
(r
1a
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7l
)e
,s
n
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o
e
m
.r
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nt
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.&
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.
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r
e
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l
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)
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l
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r i
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g.
l D
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p
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me
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o
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a
l
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g
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o
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. s
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r
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a
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l
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e
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e
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r
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t
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)
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o
n
s
.
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e
r
i
d
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s
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a
c
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e
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u
hl
m
.
(1
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ns
.
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n
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l
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d
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g
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o
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O
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e
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u
s
s
.
(
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7
8
9
)
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n
o
m
.
c
o
n
s
.
$
$
T
e
t
r
a
combi
aned
rclad
pe is
awell
echar
aacter
ized
c
and
e
Pter
a
osteemon
N
acea
ae are
kmon
aogen
ieric.
(The
1limit
9s of
4Halo
3r)agac
eae
are
h draw
en
li narr
m owly
it as
s the
o inclu
f sion
It of
e Pent
a hora
c ceae
e and
a Tetra
e carp
ar aeac
e eae
b woul
r d
o resul
a t in a
d fami
e ly
n with
e no
d obvi
b ous
e char
c acter
a s and
u totall
s y
e nove
th l
e limit
c s.
The
three
famil
ies
are
indiv
idual
ly
tolerabl
y
well
char
acter
ized.
Rece
nt
mole
cular
anal
yses
stron
gly
supp
ort a
place
ment
of
Perid
iscac
eae
withi
n
Saxif
ragal
es,
as
sister
to all
other
mem
bers
of
this
clade
(Solt
is et
al.,
2007
a;
Jian
et
al.,
2008
),
rathe
r
than
in
Malp
ighia
le Wur
s dack
a&
s Davi
p s,
re 2009
vi ),
o but
u all
sl mem
y bers
p of
r the
o expa
p nded
o fami
s ly
e have
d. simil
P ar
er disid tincti
is ve
c seed
a s etc.
c
e
a
e C
c yno
o mori
nt acea
in e are
u anot
e her
to fami
b ly of
e holo
e para
x sitic
p angi
a ospe
n rms
d that
e have
d been
( diffi
D cult
a to
vi plac
s e.
& Som
Ce
h mole
a cular
s anal
e, yses
2 had
0 plac
0 ed
4; them
in
Sant
alale
s
(Jian
et
al.,
2008
),
altho
ugh
with
little
supp
ort.
How
ever,
Bark
man
et al.
(200
7)
foun
d no
supp
ort
for a
posit
ion
in
that
order
or
any
wher
e
else.
Neve
rtheless,
Nick
rent
(200
2)
and
Nick
rent
et al.
(200
5)
sugg
ested
that
Cyn
omor
iacea
e
shou
ld be
place
d in
S stron
a g.
xi Conf
fr ound
a ing
g the
al plac
e eme
s, nt of
b this
ut taxo
th n is
e evid
e ence
vi for
d horiz
e ontal
n gene
c trans
e fer
f invol
o ving
r its
pl host
a for
ci som
ne
g mito
th chon
em drial
h gene
er s
e (Bar
v kma
er n et
s al.,
u 2007
s ).
in Zhan
S g, Li
a & Li
nt (200
al 9)
al anal
e yzed
s sequ
is ence
ns
ot from
e
Li
n
n
2
ea
0
n
0
S
9
o
T
ci
h
et
the
plast
id
inver
ted
repe
at
and
foun
d
that
Cyn
omo
rium
fell
as
sister
to
Rosa
ceae
(Ros
ales)
with
high
boot
strap
supp
ort
(99
%).
Due
to
these
disco
rdant
resul
ts,
we
do
not
assig
n
Cyn
omor
iacea
e to
an
order
here.
y anical
of Journal
L of the
o Linnea
n n
d Society,
o 2009,
n, 161,
B 105
ot 121
112
Oxalidales because
a number of recent
studies
(e.g.
Wurdack & Davis,
2009)
have
indicated that they
are
sister
to
Oxalidales
as
recognized
in
previous versions
of APG. This is not
a
wellcharacterized
clade,
and
it
remains
poorly
understood.
Malpighiale
s Juss. ex
Bercht.
& J.Presl
(1820)
Achariac
eae
Harms
(1897),
nom.
cons.
Balanopaceae
Benth. &
Hook.f. (1880),
nom. cons.
Bonnetiaceae
L.Beauvis. ex
Nakai (1948)
*Calophyllacea
e J.Agardh
Caryocaraceae
Voigt (1845),
nom. cons.
*Centroplacace
ae Doweld &
Reveal (2005) $
$Chrysobalanac
eae R.Br.
(1818), nom.
cons.
Clusiaceae
Lindl. (1836),
nom. cons.
Ctenolophonac
eae Exell &
Mendona
(1951) $
$Dichapetalace
ae Baill.
(1886), nom.
cons.
Elatinaceae
Dumort.
(1829), nom.
cons. $
$Erythroxylac
eae Kunth
(1822), nom.
cons.
(
in
cl
ud
in
g
A
n
e
ul
o
p
h
us
B
en
th.
)
E
up
ho
rb
ia
ce
ae
Ju
ss
.
(1
78
9)
,
no
m
.
co
ns
.$
$
E
up
hr
on
ia
ce
ae
M
ar
c.
B
er
ti
(1
98
9)
G
ou
pi
ac
ea
e
M
ie
rs
(1
86
2)
Humiriaceae
A.Juss. (1829),
nom. cons.
Hypericaceae
Juss. (1789),
nom. cons.
Irvingiaceae
Exell &
Mendona
(1951), nom.
cons.
Ixonanthaceae
Planch. ex Miq.
(1858), nom.
cons.
Lacistemataceae
Mart. (1826),
nom. cons.
Linaceae DC. ex
Perleb (1818),
nom. cons.
Lophopyxidacea
e H.Pfeiff.
(1951)
Meisn. (1842)
*Raffl
esiace
ae
Dumo
rt.
(1829
),
nom.
cons.
$
$Rhiz
ophor
aceae
Pers.
(1807
),
nom.
cons.
Salica
ceae
Mirb.
(1815
),
nom.
cons.
$
$Trig
oniac
eae
A.Jus
s.
(1849
),
nom.
cons.
Viola
ceae
Batsc
h
(1802
),
nom.
cons.
Malpighiaceae
Juss. (1789),
nom. cons.
$Ochnaceae DC.
(1811), nom.
cons. (including
Medusagynac
eae Engl. &
Gilg, nom.
cons.,
Quiinaceae
Choisy, nom.
cons.)
Pandaceae Engl.
& Gilg (1912
1913), nom.
cons.
$Passifloraceae
Juss. ex Roussel
(1806), nom.
cons. [including
Malesherbiaceae
D.Don, nom.
cons.,
Turneraceae
Kunth ex DC.
(1828), nom.
cons.]
Phyllanthaceae
Martinov
(1820), nom.
cons.
Picrodendracea
e Small (1917),
nom. cons.
Podostemaceae
The
Rich. ex Kunth
holoparasitic
(1816), nom.
Rafflesiaceae are
cons.
Putranjivaceae best assigned to
Linnean
2009 The Society of
London,
Malpighiales,
perhaps making
Euphorbiaceae
s.s. paraphyletic
(e.g. Davis &
Wurdack, 2004;
Davis
et
al.,
2007);
the
recognition
of
Peraceae Klotzsch
(1859) would be
needed
to
maintain
monophyly
of
Euphorbiaceae.
However, pending
further
studies,
Peraceae are not
recognized here.
Limits of clades
in
the
Bonnetiaceae
Podostemaceae
area are becoming
clearer (Wurdack
& Davis, 2009),
and this necessitates the removal
of Calophyllaceae
from Clusiaceae.
The alternatives
would be a family
that included both
of these families
and Bonnetiaceae,
Hypericaceae and
Podostemaceae or
one that included
the
last
two
families
plus
Calophyllaceae;
in both cases
Hyperi-caceae
would be the
correct name. The
four families in
the
area
of
Chrysobalanaceae
, Dichapetalaceae,
a pend
n ing a
d more
le detai
ss led
s samp
o ling
p of
h the
yl gene
o ra.
g Tric
e host
n epha
et nus
ic Gilg,
al unpl
ly aced
( previ
C ously
h,
asshare
e s
et woo
ad
l., anat
2 omy,
0 disc
0 lobin
2)g
, and
re seed
c struc
o ture
g with
ni Sam
- ydea
ti e
o Vent.
n (=
ofSalic
m acea
ore),
e and
fa unpu
m blish
ilied
esDNA
in data
th supp
is ort
ar this
e place
a ment
is (M.
prAlfo
e rd,
m pers.
at com
urm.).
e Rhiz
opho
racea
e are
kept
separ
ate
from
their
sister
taxo
n,
Eryt
hrox
ylace
ae,
altho
ugh
Aneu
loph
us,
of
Eryt
hrox
ylace
ae, is
to a
certa
in
exte
nt
morp
holo
gically
inter
medi
ate;
the
two
famil
ies
have
hithe
rto
not
been
com
bine
d.
Passi
flora
ceae
and
Och
nace
ae
are
broa
dly
deli
mite
d wo
h famil
er ies
e are
y unce
et rtain,
re and
m both
ai Med
n usag
re ynac
a eae
di (Och
ly nace
c ae
h s.l.)
ar and
a Male
ct sher
er bi- acea
iz e
a (Pass
bl iflor
e; acea
re e
la s.l.)
ti are
o mon
n ogen
s eric.
hi
p
s
b
et
C
w
u
e
c
e
u
n
r
th
b
e
i
c
t
o
a
m
l
p
e
o
s
n
e
J
nt
u
cl
s
a
s
d
.
es
w
e
it
x
hi
n
B
th
e
e
r
t
c
h
t
.
&
J
.
P
r
e
s
l
(
1
8
2
0
)
A
n
i
s
o
p
h
y
l
l
e
a
c
e
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e
R
i
d
l
.
(
1
9
2
2
)
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e
g
o
n
i
a
c
e
a
e
r
y
n
o
c
a
r
p
a
c
( e
1 a
8 e
2
4 E
) n
, g
n l
o .
m
. (
c 1
o 8
n 9
s 7
. )
C ,
o
r n
i o
a m
r .
i
a c
c o
e n
a s
e .
C
.
A
g
a
r
d
h
D
C
.
(
1
8
2
4
)
,
n
o
m
.
c
o
n
s
.
C
o
C
u
c
u
r
b
i
t
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c
e
a
e
J
u
s
s
.
(
1
7
8
9
)
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n
o
m
.
c
o
n
s
.
D
a
t
i
s
c
a
c
e
a
e
D
u
m
o
r
t
.
(
1
8
2
9
)
,
n
o
m
.
c
o
n
s
.
T
e
t
r
a
m
e
l
a
c
e
a
e
A
i
r
y
S
h
a
w
(
1
9
6
5
)
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a
b
al
es
B
ro
m
h
e
a
d
(1
8
3
8)
F
a
b
a
c
e
a
e
,
no
m.
co
ns.
Po
ly
ga
la
ce
ae
H
of
fm
an
ns.
&
Li
nk
(1
80
9),
no
m.
co
ns.
Q
uil
laj
ac
ea
e
D.
D
on
(1
83
1)
Su
ria
na
ce
ae
Ar
n.
(1
83
4),
no
m.
co
ns.
LFaga
iles
nEngl.
d(189
l2)
. B
(
e
1
8 t
3 u
6 l
) a
c
e
a
e
G
r
a
y
(
1
8
2
2
)
,
n
o
m
.
c
o
n
s
.
C
a
s
u
a
r
i
n
a
c
e
a
e
R
.
B
r
.
(
1
8
1
4
)
,
n
o
m
.
c
o
n
s
.
F
a
g
a
c
e
a
e
D
u
m
o
r
t
.
(
1
8
2
9
)
,
n
o
m
.
c
o
n
s
.
Ju
g
l
a
n
d
a
c
e
a
e
D
C
.
e
x
P
e
r
l
e
b
z
.
(
1
8
1
8
)
,
(
1
9
3
2
)
,
n
o
m
.
n
o
m
.
c
o
n
s
.
c
o
n
s
.
]
[
i
n
c
l
u
d
i
n
g
R
h
o
i
p
t
e
l
e
a
c
e
a
e
H
a
n
d
.
M
a
z
M
y
r
i
c
a
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A
.
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(
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1
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)
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o L.
m D.
. G
c
o m
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n
(1
s 99
. 1)
N
o Ju
tglan
hdace
oae
fare
aexpa
gnded
ato
c
inclu
e
de
a
their
e
sister
clade
K
u, the
pmon
rogen
ieric
aRhoi
nptele
acea
(e.
1The
9
two
6
look
2
)
T
i
c
o
d
e
n
d
r
a
c
e
a
e
G
m
e
z
L
a
u
r
.
&
simil
ar,
even
altho
ugh
Rhoi
ptele
acea
e
have
supe
rior
and
Jugla
ndac
eae
infer
ior
ovari
es;
ovar
y
posit
ion
is
varia
ble
in
man
y
famil
ies,
and
in
other
such
cases
, e.g.
Eben
acea
e
(incl
udin
g
Liss
ocar
pace
ae),
we
have
reco
gniz
ed
the
large
r
unit.
R
o
s
a
l
e
s
o
m
.
c
o
n
Bs
e.
r
cC
ha
t n
. n
a
&b
a
J c
. e
Pa
r e
e
s M
l a
r
( t
1i
8n
2o
0v
)
(
B1
a8
r 2
b0
e)
y,
a
cn
eo
am
e.
Rc
eo
nn
ds
l .
e
D
( i
1r
9a
1c
6h
) m
, a
c
ne
a
e
H
u
t
c
h
.
(
1
9
5
9
)
E
l
a
e
a
g
n
a
c
e
a
e
J
u
s
s
.
(
1
7
8
9
)
,
n
o
m
.
c
o
n
s
.
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o
r
a
c
e.
a
ec
o
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as
u.
d
i R
co
hs
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c
( e
1a
8e
3
5J
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o
m(
. 1
7
c8
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s ,
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n
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ao
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e
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J l
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s c
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a
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1
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n(
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5
)
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m
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c
o
n
s
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U
r
t
i
c
a
c
e
a
e
J
u
s
s
.
(
1
7
8
9
)
,
n
o
m
.
c
o
n
s
.
MALVIDS
Gera
ni
al
es
Ju
s
s.
e
x
B
e
r
c
h
t.
&
J
.
P
r
e
sl
(
1
8
2 $
0
)
$
G
e
r
a
n
ia
c
e
a
e
J
u
s
s.
(
1
7
8
9
),
n
o
m
.
c
o
n
s.
(i
n
cl
u
d
i
n
g
Hy
p
s
e
o
c
h
a
r
i
t
a
c
e
a
e
(
i
n
c
l
u
d
i
n
g
F
r
a
n
c
o
a
c
e
a
e
W
e
d
d
.
)
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e
l
i
a
n
t
h
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c
e
a
e
A
.
J
u
s
s
.
,
n
o
m
.
H
o
r
a
n
.
(
1
8
3
4
)
,
n
o
m
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c
o
n
s
.
c
o
n
s
.
)
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i
v
i
a
n
i
a
c
e
a
e
K
l
oe a
theter
zogen
seous
cand
hpoorl
,y
kno
n
wn
o
order
m
. The
.
inclu
csion
oof
nthe
smon
.ogen
peric
rHyps
oeoch
pari.tacea
(e in
iGera
nniace
cae,
mon
l
ogen
u
eric
d
Fran
i
coac
neae
gin
LMeli
eanth
dacea
oe and
cbige
aneric
rLedo
pcarp
aacea
ce in
eVivi
aaniac
eae
e
leave
M
s
e
these
y
expa
ended
nfamil
)ies
with
er a
a num
ni ber
al of
eschar
ar acter
s.
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y
r
t
a
l
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s
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u
s
s
.
e
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e
r
c
h
t
.
&
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.
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r
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l
(
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)
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l
z
a
t
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a
c
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.
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r a
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me
(
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)
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C
.
(
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r )
e,
t
an
co
em
a.
e
c
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. n
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r .
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( y
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8h
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0a
) c
, e
a
ne
o
mJ
. .
S
ct
o.
ns H
. i
l
C.
r
y(
p1
t 8
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r 5
o)
n,
n
o
m
.
c
o
n
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.
$
M
e
l
a
s
t
o
m
a
t
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c
e
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e
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u
s
s
.
(
1
7
8
9
)
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o
m
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c
o
n
s
.
(
i
n
c
l
u
d
-
i
n
g
M
e
m
e
c
y
l
a
c
e
a
e
D
C
.,
n
o
m
.
c
o
n
s
.
)
M
y
r
t
a
c
e
a
e
J
u
s
s
.
(
1
7
8
9
)
,
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o
m
.
c
o
n
s
.
(
i
n
c
l
u
d
i
n
g
H
e
t
e
r
o
p
y
x
i
d
a
c
e
a
e
E
n
g
l
.
&
G
i
l
g
,
n
o
m
.
c
o
n
s
.
,
P
s
i
l
o
x
y
l
a
c
e
a
e
C
r
o
i
z
a
t
)
O
na
gr
ac
ea
e
Ju
ss.
(1
78
9)
,
no
m.
co
ns
.
P
e
n
a
e
a
c
e
a
e
S
w
e
e
t
e
x
G
u
i
l
l
.
(
1
8
2
8
)
,
L
n
.
o
A
m .
.
S
c
.
o
J
n
o
s
h
.
n
(
s
i
o
n
n
c
l
&
u
d
B
i
.
n
G
g
.
O
B
l
r
i
i
n
g
i
g
a
s
c
)
e V
a oc
e hy
A si
r ac
ea
n
e
., A.
n St
o .m Hi
. l.
c (1
o 82
n 0)
s ,
no
.,
m.
R
co
h ns
y .
n
- In
cMela
hstom
oatace
cae,
aand
lstill
ymore
cin
aMyrt
cacea
ee,
acom
emon
usag
e is
for
broa
dene
d
famil
y
circu
mscr
iptions
.
Both
Hete
ropy
xida
ceae
and
Psilo
xyla
ceae
are
smal
l
famil
ies
and
whe
n
inclu
ded
in
Myrt
acea
e s.l.
that
famil
y
rema
ins
char
acter
ized
by
poss
essio
n of
pellu
cid
glan
ds
cont
ainin
g
ether
eal
oils.
A
close
relati
onsh
ip rypte
b ronia
et ceae,
w Pena
e eace
e ae,
n Olini
C acea
e
Li
n
2n
0 ea
0n
9S
To
h ci
et
e,
Alza
taeac
eae
and
Rhy
ny anical
of Journal
L of the
o Linnea
n n
d Society,
o 2009,
n, 161,
B 105
ot 121
114
supported,
andBrassicales
recognition
ofBromhead (1838)
Huerteales
is $Akaniaceae
Stapf (1912),
appro-priate given
nom. cons.
their
position.
(including
Within Huerteales,
Bretschneide
the
recently
raceae Engl.
described
& Gilg, nom.
Gerrardinaceae
cons.)
(Gerrardina was
Batacea
previously placed
in Flacourtiaceae) e Mart.
are sister to the ex
rest,
and Perleb
Dipentodontaceae, (1838),
nom.
although
monogeneric like cons.
Gerrardinaceae, are Brassic
distinctive
(see aceae
Worberg et al., Burnett
(1835),
2009).
nom.
cons.
*Cappar
aceae
Juss.
(1789),
nom.
cons.
Caricac
eae
Dumort.
(1829),
nom.
cons.
*Cleom
aceae
Bercht.
&
J.Presl
(1825)
Embling
iaceae
J.Agard
h (1958)
Gyroste
monace
ae
A.Juss.
(1845),
nom.
cons.
Koeberl
iniaceae
Engl.
(1895),
nom.
cons.
Limnant
haceae
R.Br.
(1833),
nom.
cons.
Moringa
ceae
Martino
v
(1820),
nom.
cons.
Pentadi
plandrac
eae
Hutch.
&
Dalziel
(1928)
Resedac
eae
Martino
v
(1820),
nom.
cons.
Salvado
raceae
Lindl.
(1836),
nom.
cons.
Setchell
anthace
ae Iltis
(1999)
Tovariace
ae Pax
(1891),
nom.
cons.
Tropaeola
ceae Juss.
ex DC.
(1824),
nom.
cons.
Inclusion
of
monogeneric
Bretschneideracea
e
into
the
monogeneric
Akaniaceae
is
justified by the
morphological
similarities of the
two, which are
sister
taxa.
Although a broad
circumscription of
Brassicaceae was
recognized in APG
(1998) and APG II
(2003),
the
consensus prefers
the recognition of
three families, all
of which can be
characterized,
albeit Capparaceae
only rather poorly
so.
The
final
phylogenetic
positions,
and
hence taxonomic
disposi-tion,
of
some
genera,
particularly those
previously
included
in
Capparaceae
Stixeae,
remain
uncertain
(Hall,
Sytsma & Iltis,
2002; Hall, Iltis &
Sytsma,
2004).
Nonetheless,
the
name
Stixaceae
Doweld (2008) is
available if it is
required.
Malvales Juss. ex
Bercht. & J.Presl
(1820)
$Bixaceae
Kunth (1822),
nom.
cons.
(including
Cochlosperma
ceae Planch.,
nom. cons.,
DiegodenLinnean
2009 The Society of
London,
draceae
Capuron,)
Cistac
eae
Juss.
(1789),
nom.
cons.
*Cytin
aceae
A.Rich
.
(1824)
Dipter
ocarpa
ceae
Blume
(1825),
nom.
cons.
Malva
ceae
Juss.
(1789),
nom.
cons.
Muntingiaceae
C.Bayer,
M.W.Chase
& M.F.Fay
(1998)
Neurad
aceae
Kostel.
(1835),
nom.
cons.
Sarcola
enacea
e
Caruel
(1881),
nom.
cons.
Sphaer
osepala
ceae
Tiegh.
ex
Bulloc
k
(1959)
Thymel
aeaceae
Juss.
(1789),
nom.
cons.
A
broad
circumscription
for Bixaceae is
adopted; the three
families included
are all small, and
the
combined
family can be
characterized
morphologically.
The
para-sitic
Cytinaceae
(including
Bdallophyton
Eichl.) find their
resting place here
(Nickrent, 2007).
The
novel
dismemberment
of Malvaceae by
Cheek (2006), see
also Cheek in
Heywood et al.,
2007)
is
not
followed
h the
Sarc
er first
olae
e; is a
later
nace
th
nam
ae
e
e
for
Dipt
fa
Spar
eroc
m
man
arpa
ili
ceae
esniace
rema
ar ae
in
e J.Ag
ardh
uncl
di
as
ear,
ff
defin
and
ic
ed
these
ul
by
famil
t
ies
to Chee
may
di k).
need
st The
close
to be
in
relati
com
g
onsh
bine
ui
ip
of
d
s
the
(Kub
h,
itzki
a four
&
n famil
Chas
d ies
that
e,
t
mak
2002
w
e
up
;
o
Malv
Duc
ar
acea
ouss
e
o et
n e s.l.
al.,
e here
2004
w has
been
);
(
reco
Cista
B
gniz
ceae
ro
ed
has
w
since
prior
nl
ity if
o at
these
w least
are
ia the
time
all
c
of
com
e
Robe
bine
a
rt
d as
e,
Bro
a
D
singl
urwn.
e
io Deta
famil
n ils of
relati
y.
a
onsh
c
e ips
Sa
a in
p
e, the
i
al area
n
th of
d
o Cista
a
l
e
s
1
8
1
8
)
J ,
u
s n
s o
. m
.
e
xc
o
Bn
es
r .
c
hB
t i
. e
b
&e
r
J s
. t
Pe
r i
en
s i
l a
c
( e
1a
8e
2
0S
) c
h
An
ni
az
cl
a.
r
d(
i 1
a8
c5
e6
a)
e
B
Ru
. r
Bs
r e
. r
a
( c
e
a
e
K
u
n
t
h
(
1
8
2
4
)
,
n
o
m
.
c
o
n
s
.
K
i
r
k
i
a
c
e
a
e
T
a
k
h
t
.
(
1
9
6
7
)
M
eli
ac
ea
e
Ju
s
s
.
(
1
7
8
9
)
,
n
o
m
.
c
o
n
s
.
$N
i
t
r
a
r
i
a
c
e
a
e
L
i
n
d
l
.
(
1
8
3
5
)
,
n
o
m
.
c
o
n
s
.
(
i
n
c
l
u
d
i
n
g
P
e
g
a
n
a
c
e
a
e
T
i
e
g
h
.
e
x
T
a
k
h
t
.
,
T
e
t
r
a
d
i
c
l
i
d
a
c
e
a
e
T
a
k
h
t
.
)
R
u
t
a
c
e
a
e
J
u
s
s
.
(
1
7
8
9
)
,
n
o
m
.
c
o
n
s
.
S
a
p
i
n
d
a
c
e
a
e
J
u
s
s
.
(
1
7
8
9
)ceae
,broa
ndly.
oThe
m
four
.gene
cra
oinclu
nded
sshow
.consi
Sdera
ible
m
varia
ation,
raltho
ough
utheir
bbasic
amorp
cholo
egy,
aanat
eomy
and
D
che
C
m.istry
(are
1poorl
8y
1kno
1wn.
)
,
nBer
berid
oopsi
m
dales
.Dow
celd
o(200
n1)
s A
. ex
to
xi
e ca
ci ce
ae
rc
En
u gl.
m &
sc Gi
ri lg
b (1
e 92
0),
N
no
it m.
ra co
ri ns.
a Be
rb
eri
do
ps
id
ac
ea
e
Ta
kh
t.
(1
98
5)
Th
e
morp
holo
gical
ly
disti
nct
Aext
oxic
acea
e and
Berb
erido
psida
ceae
are
stron
gly
supp
orted
as
sister
taxa,
and
rece
nt
work
(Mo
ore
et
al.,
in
press
)
place
d
them
with
stron
g
supp
ort
as
sister
to
(Sant
al
al B
ese
( r
Cc
ar h
yt
o.
p
h&
yl
la J
le .
s P
+r
ase
te s
ri l
d
s) (
); 1
th 8
u2
0
s,
)
or
di
*
n
B
al
a
st
l
at
a
u
n
s
o
is
p
a
h
p
o
prr
oa
prc
ia e
te a
. e
Sa R
ni
t c
ah
l .
a
l (
e1
s 8
2
R2
. )
B,
r
. n
o
em
x.
c
o
n
s
.
L
o
r
a
n
t
h
a
c
e
a
e
J
u
s
s
.
(
1
8
0
8
)
,
n
o
m
.
c
o
n
s
.
M
i
s
o
d
e
n
d
r
a
c
e
a
e
J
.
A
g
a
r
d
h
(
1
8
5
8
)
,
n
o
m
.
c
o
n
s
.
S
a
n
t
a
l
a
c
e
a
e
R
.
B
r
.
(
1
8
1
0
)
,
n
o
m
.
c
o
n
s
.
O
l
a
c
a
c
e
a
e
R
.
B
r
.
(
1
8
1
8
)
,
n
o
m
.
c
o
n
s
.
O
p
i
l
i
a
c
e
a
e
V
a
l
e
t
o
n
(
1
8
8
6
)
,
n
o
m
.
c
o
n
s
.
*
S
c
h
o
e
p
f
i
a
c
e
a
e
B
l
u
m
e
(
1
8
5
0
)
Th
e
gene
ra
inclu
ded
in
Scho
epfia
ceae
used
to be
inclu
ded
in
Olac
acea
e s.l.,
but
they
are
excl
u morp
si holo
v giel cally
y so
re disti
la nct
te that
d. com
T binat
h ion
e of
y the
ar two
e famil
w ies is
el inap
l prop
s riate.
u The
p para
p phyl
oretic
te Olac
d acea
ase are
b bein
ei g
n resol
g ved
in into
a a
cl num
a ber
d of
e clade
ws
it (Mal
h cot
M&
is Nick
o rent,
d 2008
e ), but
n relati
dronsh
a ips
c betw
e een
a these
e, clade
b s are
ut unce
th rtain
at and
fa so
m new
il famil
y ies/f
is amil
y
limit
s
ar ded
e into
n clade
ot s
pr(Der
o&
p Nick
o rent,
se2008
d ),
h one
er of
e. whic
S h is
a the
nt morp
al holo
a gical
c ly
e disti
a nct
e Visc
ar acea
e e, is
k of
e itself
pt insuf
w ficie
it nt
h reaso
th n for
ei their
r dism
premb
e erme
vi nt
o (see
u Intro
s ducti
ci on,
rc also
u Dips
m acale
scs
ri belo
pt w).
io Bala
n. noph
T oh racea
at e are
th to be
e inclu
y ded
c in
a Sant
n alale
bs
e (Nic
di krent
vi , Der
&
And
erso
n,
2005
;
Bark
man
et
al.,
2007
),
and
there
is
some
evid
ence
that
Cyn
omor
iacea
e
migh
t also
belo
ng
here
(see
com
ment
s
unde
r
Saxif
ragal
es,
abov
e).
Cary
op
hy
lla
le
s
Ju
ss.
ex
B
er
ch
t.
&
J.
Pr
es
l
(1
82
0
)
A
c
h
at
o
c
a
r
p
a
c
e
a
e
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ei
m
e
rl
(
1
9
3
4
),
n
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m
.
c
o
n
s.
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iz
o
a
c
e
a
e
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a
rt
i
n
o
v
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1
8
2
0
),
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o
m
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co
ns
.
A
m
ar
an
th
ac
ea
e
Ju
ss.
(1
78
9)
,
no
m.
co
ns
.
*
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na
ca
m
ps
er
ot
ac
ea
e
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gg
li
&
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yf
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le
r
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01
0,
in
p
r
e
s
s
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n
c
i
s
t
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l
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h
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a
l
p
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1
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o
m
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c
o
n
s
.
A
st
er
op
ei
ac
ea
e
Ta
kh
t.
ex
R
ev
ea
l
& n
o
H m
o .
o
g c
l o
a n
n s
d .
(
1 C
9 a
9 c
0 t
) a
B c
a e
r a
b e
e
u J
i u
a s
c s
e .
a
e (
N 1
a 7
k 8
a 9
i )
( ,
1
9 n
4 o
2 m
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a c
s o
e n
l s
l .
a
c C
e a
a r
e y
R o
a p
f h
. y
( l
1 l
8 a
3 c
7 e
) a
, e
J
u
s
s
.
(
1
7
8
9
)
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o
m
.
c
o
n
s
.
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i
d
i
e
r
e
a
c
e
a
e
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a
d
l
k
.
(
1
8
9
6
)
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n
o
m
.
c
o
n
s
.
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i
o
n
c
o
p
h
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l
l
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e
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i
r
y
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h
a
w
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1
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o
n
s
.
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r
o
s
e
r
a
c
e
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e
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a
l
i
s
b.
(1
80
8)
,
no
m.
co
ns
.
D
ro
so
ph
yl
la
ce
ae
C
hr
te
k,
Sl
av
k
ov
&
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ud
ni
c
ka
(
1
9
8
9
)
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r
a
n
k
e
n
i
a
c
e
a
e
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e
s
v
.
(
1
8
1
7
)
,
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o
m
.
c
o
n
s
.
G
i
s
e
k
i
a
c
e
a
e
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a
k
a
i
(
1
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H
a
l
o
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h
y
t
a
c
e
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e
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.
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o
r
i
a
n
o
(
1
9
8
4
)
*
L
i
m
e
a
c
e
a
e
S
h
i
p
u
n
o
v
e
x
R
e
v
e
a
l
(
2
0
0
5
)
*
L
o
p
h
i
o
c
a
r
p
a
c
e
a
e
D
o
w
e
l
d
&
R
e
v
e
a
l
(
2
0
0
8
)
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o
l
l
u
g
i
n
a
c
e
a
e
B
a
r
t
l
.
(
1
8
2
5
)
,
n
o
m
.
c
o
n
s
.
*
M
o
n
t
i
a
c
e
a
e
R
a
f
.
(
1
8
2
0
)
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e
p
e
n
t
h
a
c
e
a
e
D
u
m
o
r
t
.
(
1
8
2
9
)
,
n
o
m
.
c
o
n
s
.
N
y
c
t
a
g
i
n
a
c
e
a
e
J
u
s
s
.
(
1
7
8
9
)
,
n
o
m
.
c
o
n
s
.
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h
y
s
e
n
a
c
e
a
e
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a
k
h
t.
(
1
9
8
5
)
P
h
y
t
o
l
a
c
c
a
c
e
a
e
R
.
B
r
.
(
1
8
1
8
)
,
n
o
m
.
c
o
n
s
.
P
l
u
m
b
a
g
i
n
a
c
e
a
e
J
u
s
s
.
(
1
7
8
9
)
,
n
o
m
.
c
o
n
s
.
P
o
l
y
g
o
n
a
c
e
a
e
J
u
s
s
.
(
1
7
8
9
)
,
n
o
m
.
c
o
n
s
.
P
o
r
t
u
l
a
c
a
c
e
a
e
J
u
s
s
.
(
1
7
8
9
)
,
n
o
m
.
c
o
n
s
.
R
h
a
b
d
o
d
e
n
d
r
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a
e
P
r
a
n
c
e
(
1
9
6
8
)
S
a
r
c
o
b
a
t
a
c
e
a
e
m
a
t
a
c
e
a
e
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e
h
n
k
e
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a
k
a
i
(
1
9
9
7
)
(
1
9
4
2
)
S
i
m
m
o
n
d
s
i
a
c
e
a
e
*
T
a
l
i
n
a
c
e
a
e
T
i
e
g
h
.
(
1
9
0
0
)
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t
e
g
n
o
s
p
e
r
D
o
w
e
l
d
(
2
0
0
1
)
Ta
m
ar
ic
ac
ea
e
Li
nk
(1
82
1)
,
nmilie
os is
m
nece
.ssitat
c
ed
o
nby
srece
.nt
phyl
ogen
h etic
e work
re on
c core
o Cary
g ophy
ni llales
ti .
o Anac
n amps
oferota
a ceae,
n Port
u ulam cace
b ae
er s.s.,
ofMon
n tiace
e ae
w and
b Talin
ut acea
s e are
m all
al clade
l s
fa near
e
Li
n
2n
0 ea
0n
9S
To
h ci
et
Cact
acea
e
that
are
for
the
most
part
well
supp
orted
as
disti
nct
(e.g.
Appl
equis
t &
Wall
ace,
2001
;
Nyff
eler,
2007
;
Nyff
eler
&
Eggl
i, in
press
;
Broc
kingt
on
y anical
of Journal
L of the
o Linnea
n n
d Society,
o 2009,
n, 161,
B 105
ot 121
116
cons.
Roridulac
eae
Martinov
(1820),
nom. cons.
Sapotacea
e Juss.
(1789),
nom. cons.
Sarracenia
ceae
Dumort.
(1829),
nom. cons.
$
$Sladenia
ceae Airy
Shaw
(1965)
Styracacea
e DC. &
Spreng.
(1821),
nom. cons.
Symploca
ceae Desf.
(1820),
nom. cons.
Marcgraviaceae
Bercht. &
J.Presl (1820),
nom. cons.
*Mitrastemonac
eae Makino
(1911), nom.
cons.
$Pentaphylacace
ae Engl. (1897),
nom. cons.
(including
Ternstroemiacea
e Mirb. ex DC.)
Polemoniaceae
Juss. (1789),
nom. cons.
Primulaceae
Batsch ex
Borkh. (1797),
nom. cons.
(including
Maesaceae
Anderb.,
B.Sthl
&
Kllersj,
$Tetrameristace
Myrsinaceae
ae Hutch.
R.Br., nom.
(1959)
cons., Theo(including
phrastaceae
Pellicieracea
G.Don, nom.
e L.Beauvis.)
cons.)
Theaceae Mirb.
ex Ker Gawl.
(1816), nom.
cons.
It was clear in
APG
II
that
Theaceae
s.l.
could
not
be
maintained.
Subsequent work
on the potential
segregates
has
clarified
the
morphological
pattern
of
variation
(Stevens,
2001,
for a summary).
Sladeni-aceae are
recognized
as
distinct
from
Pentaphylacaceae; although
the two are sister
taxa, they share
few
obvious
characters,
and
little would be
gained by uniting
them. However,
Ternstroemiaceae
have much in
common
with
Pentaphylacaceae
and so the former
are included in the
latter. Theaceae
s.s. are not immediately related to
these families.
The
monogeneric
Pellicieraceae are
included
in
Tetrameristaceae;
the
resulting
family, with three
genera,
is
moderately well
characterized.
Mitrastemonaceae
is
a
morphologically
distinctive
holoparasitic
family that is well
embedded
in
Ericales.
The
biggest
problem for APG
III
was
the
question of how
to
treat
Primulaceae and
their immediate
relatives, a closely
related group that
in the past has
often
been
recognized as a
separate
order.
Although
Primulaceae and
relatives
are
clearly
in
Ericales,
taxon
limits in this
group have been
problematic.
Maesaceae are a
monogeneric
family
necessitated
by
the break-up of
Myrsinaceae, as
APG III
Metteniusaceae H.Karst. ex Schnizl. (18601870)
Oncothecaceae Kobuski ex Airy Shaw (1965)
The limits
of
Boraginaceae
are drawn broadly.
Not only are
the
phylogenetic relationships
within
the family still
unclear, but as we know
more
about relationships within its component clades, they
become less easy to distinguish (e.g. Gottschling et al.,
2005 for Cordioideae A.Gray). Molecular data suggest
that Hoplestigmataceae are to be included in Boraginaceae s.l., being placed in or near Cordioideae (K.
Wurdack, pers. comm.; V. Savolainen and M. Powell,
pers. comm.); Hoplestigma Pierre is similar in inflorescence, ovary, pollen, etc. to Boraginaceae. Relationships of Boraginaceae s.l. and Vahliaceae remain
117
CAMPANULIDS
Aquifoliales Senft (1856)
Aquifoliaceae Bercht. & J.Presl (1820), nom. cons.
Cardiopteridaceae Blume (1847), nom.
(including Leptaulaceae Tiegh.)
Helwingiaceae Decne. (1836)
Phyllonomaceae Small (1905)
Stemonuraceae Krehed (2001)
Leptaulus Benth., previously unplaced, is assigned
to Cardiopteridaceae (Krehed, 2001).
Asterales Link (1829)
Alseuosmiaceae Airy Shaw (1965)
Argophyllaceae Takht. (1987)
Asteraceae Bercht. & J.Presl (1820), nom. cons.
Calyceraceae R.Br. ex Rich. (1820), nom. cons.
$Campanulaceae Juss. (1789), nom. cons. (including Lobeliaceae Juss., nom. cons)
Goodeniaceae R.Br. (1810), nom. cons.
Menyanthaceae Dumort. (1829), nom. cons.
Pentaphragmataceae J.Agardh (1858), nom. cons.
cons.
118
3
6
)
,
n
o
m
.
c
o
n
s
.
A
r
a
l
i
a
c
e
a
e
J
u
s
s
.
(
1
7
8
9
)
,
n
o
m
.
c
o
n
s
.
Griseliniaceae
J.R.Forst. &
G.Forst. ex
A.Cunn.
(1839)
Myodocarpacea
e Doweld
(2001)
P
e
nn
form a strongly
ant
supported clade
iac
(e.g.
Lundberg,
ea
2001;
Plunkett,
e
2001;
Krehed,
J.
2002, 2003), and
Ag
that they were
ard
kept
separate
h
before was a
(1
simple oversight.
85
The recognition of
8)
Myodocarpaceae
Pit
results from our
tos
improved
po
understanding of
rac
relationships
of
ea
members included
e
formerly
in
R.
Araliaceae.
Br.
(1
81
TAXA
4),
OF
no
UNCER
m.
TAIN
co
POSITI
ns.
ON
Torricelliaceae
Hu
(1934)Apodanthacea
e Takhtajan
(including
Aralidiaceae [three genera]
Philipson &Cynomoriacea
e Endl. ex
B.C.Stone,
Melanophylla Lindl. (1833),
ceae Takht. exnom. cons.
Airy Shaw) Gumillea Ruiz
& Pav.
Expansion
ofPetenaea Lundell
Torricelliaceae to(possibly
include
Malvales)
Aralidiaceae andNicobariodendron
Melanophyllaceae (see
Simmons,
is reasonable. All2004; possibly
three
are Celastraceae).
monogeneric and
poorly
known.
RE
Nevertheless, they
Linnean
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London,
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to
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ae
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Ja
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