Botanical Journal of the Linnean Society, 2009, 161, 105121.

With 1 figure

An update of the Angiosperm Phylogeny Group classification for

the orders and families of flowering plants: APG III
THE ANGIOSPERM PHYLOGENY GROUP*1
1

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.

ADDITIONAL KEYWORDS: angiosperm classification phylogenetic classification DNA phylogenetics classification

system APG system.

*Corresponding author: Mark W. Chase. E-mail: m.chase@kew.org

2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 161, 105121

105

106

THE ANGIOSPERM PHYLOGENY GROUP

follow
these
particular have a
principles
here.
number
of
I
Backlund
and
problems
in
this
N
main
regard and include
TBremers
principle
is
that
fami-lies that are
R
taxa
that
are
still in a state of
O
recognized
flux
in
this
D
formally
should
respect.
U
be mono-phyletic.
C
this
THowever,
Changes
are
does
not
indicate
I
being
made
to
the
which
par-ticular
O
clades should be
classification
N
named as families,
adopted by APG
Since the previousorders etc.
II for several
APG classification
reasons,
but
(APG II, 2003), a
bearing
the
great deal more It is helpful if
Backlund
and
information about(1) taxa formally
Bremer guidelines
flowering
plantrecognized
are
in mind.
relationships haseasily
emerged, and arecognizable, (2)
1. In the earlier
more refined andgroups that are
versions of the
better-resolved
Angiosperm
well estab-lished
classification
isin the literature
Phylog-eny
now possible. If aare preserved, (3)
Group
well-supported
classification,
the size of groups
hypothesis
ofis
alternative
taken
into
monophyly is aaccount
circumscripnecessary
tions
were
(particularly small
prerequisite for aones,
permitted for
which
group to be named,should
some families.
be
it is not suffi-cient.combined
However, this
with
Not all clades needothers whenever
seems
be named and,pos-sible) and (4)
unnecessary
indeed, it wouldnomenclatural
and
more
barely
bechanges
likely to cause
are
practicable to dominimized
confusion than
this, so to decide(Backlund
clarity. Here
&
which
cladesBremer,
we
indicate
1998).
should be named,Thus, on the one
one of the
additional criteriahand, numerous
alternatives, a
can be invoked.small groups have
choice based
Backlund
&little
in part on the
to
Bremer
(1998)recommend them,
circumprovided a usefulas
scriptions
individually
discussion on thethey summarize
adopted
by
principles of rank-only
textbooks (e.g.
a
small
based
phy-amount
Judd et al.,
of
logenetic
2007),
information and
classification thattend to clog the
dictionaries
is applicable at allmemory, whereas
(e.g.
levels apart fromgroups that are too
Mabberley,
species
andlarge may have
2008) and the
immediately abovefew
general
obvious
(see Albach et al.,shared traits that
literature.
2004; Entwisle &can be used to
Note that, in
Weston, 2005; Pfeilrecognize them.
preparation for
& Crisp, 2005, etc.,Asparagales and
the
third
for examples). WeLamiales
edition of The
in

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

change the status of agreement

for
families if theychange of any
have long been sort. Note that we
recognized, unless do not see the
there are other APG classificagood reasons fortion as continuing
combining them. to mutate for the
However, we have indefinite future.
taken
theGiven the amount
opportunity
toof phylogenetic
combine mono- orwork that has
oligogeneric
taken place in the
families. Most of last five years, the
the
family-level changes pro-posed
changes mentioned here are modest.
below
haveWe hope
the
resulted in the classification
families
nowbelow will be
recognized beingfound
to
be
relatively
well reasonable
and,
charac-terized.
hence, will not
However,
we need much further
realize that it is change, although
almost impos-sible we do note those
to
achieve few areas where
universal
there is particular
2009 The

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

APG III 107

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Aug
ust
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ra
plant
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).
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of anical
L Journal
o of the
n Linnean
d Society,
o 2009,
n, 161,
B 105
ot 121

THE ANGIOSPERM PHYLOGENY GROUP

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
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Oxalidales
Malpighiales
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Malvales
Brassicales
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malvids
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Crossosomatales
Myrtales
Geraniales
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Saxifragales
Dilleniaceae
Berberidopsidales
Santalales
Caryophyllales
Cornales
Ericales
Garryales
Gentianales
lamiids
Lamiales
Solanales
Boraginaceae
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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

APG III 109

no
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B m.
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.
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o
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is
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T
(e.g.
o
fChen
iet
eal.,
l2004
da, b)
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Whe
n
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a
famil
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vely
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descr
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nquis
t,
1981
), the
enlar
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sever
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chara
cters.
It
may
be
nece
ssary
to
split
off
Mau
ndia
from
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agina
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(Iles
et
al.,
2009
; S.
von
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ng &

J. case
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er singl
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er the
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m More
.), study
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N gnize
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k Alis
ai mata
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a
v
ai
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bl osavi
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e.
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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),
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r
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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)
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r
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i
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o (
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e 8
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R
.J.Ag
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r
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rgical
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ges
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kx et
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).
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Aspa
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ragac
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o Journal
n, of the
B Linnean
ot Society,
a 2009,
ni 161,
c 105
al 121

110

THE ANGIOSPERM PHYLOGENY GROUP

(2009). This will
(1804),
allow
researchers
nom.
caceae and even
to
use
a
subfamily
cons.
Hyacinthaceae
name
where
Haemodo
have few if any
previously
they
raceae
dis-tinctive
would have used
R.Br.
features.
(1810),
Agavaceae
andone of the APG II
nom.
Ruscaceae sensubracketed family
names.
cons.
APG II (2003) are
Hanguana
heterogeneous and
ceae Airy
have been divided
Shaw
into
several
C
(1965)
families in the past,
O
Philydrac
further confusing
M
eae Link
the issue. This
M
(1821),
solution at least
E
nom.
keeps the number
LI
cons.
of
difficult-toNI
Pontederi
recognize families
D
aceae
to a minimum;
S
Kunth
Amaryl-lidaceae
(1816),
s.l.
andDasypogonaceae
Dumort. (1829)
nom.
Asparagaceae s.l.
cons.
are easily differen- Relationships
tiated by umbelsof
Poales Small
with a pair ofDasypogonaceae
(1903)
enclosing bracts vs.remain
unclear.
Anarthriace
racemes or rarelyThey are not
ae
umbels, but ifparticularly
D.F.Cutler
umbels are presentdistinctive
& Airy
they contain threemorphologically
Shaw
or more bracts (e.g.and so do not
(1965)
Brodiaea,
Millawarrant
Bromeliace
and
relatives,immediate ordinal
ae Juss.
Themidaceae) andstatus, although
(1789),
lack the enclosingDasypogonales
nom. cons.
pair of bracts.Doweld
is
Centrolepid
Xanthorrhoeaceae available.
They
aceae Endl.
s.l. have been
could probably be
(1836),
combined
with
nom. cons.
maintained for similar
Cyperaceae
one reasons
of the other
s.l. Xanthorrhoeaceae
Juss.
commelinid
Asphodelaceae areorders should they
(1789),
impossible
tofall
nom. cons.
as
sister
distinguish
fromgroups.
Ecdeiocole
genera such as
aceae
Anthericum
inArecales
D.F.Cutler
Asparagaceae s.l. Bromhead (1840)
& Airy
For convenience Arecaceae
Shaw
and
better Bercht. &
(1965)
J.Presl (1820),
communication, a
Eriocaulace
nom. cons.
subfamilial
ae
classification
of Commelinal
Martinov
es Mirb.
Amaryllidaceae,
(1820),
Asparagaceae and ex Bercht.
nom. cons.
& J.Presl
Xanthorrhoeaceae
Flagellariac
sensu APG III is (1820)
eae
proposed in Chase, Commeli
Dumort.
Reveal & Fay naceae
(1829),
Mirb.

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.,

2009). In this and

all
other
relationships that
have
been
suggested
for
Ceratophyllaceae,
including sister to
the monocots or
Chloranthaceae
(Endress & Doyle,
2009), they are
morphologically
divergent
from
their
putative
closest relatives.

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)

Botanical Society, 2009, 161, 105

Journal of 121
the Linnean

APG III 111

e
broa
dene
ar dich
d
otom
e
circu
ous
b
mvena
ot
scrip
h tion.
tion
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Prote
n acea
ales;
oe
the
Blu
g
two
me
e (185
have
n 1),
featu
res
er nom.
cons.
in
ic
com
;
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mon.
th thou
Sabi
e gh
acea
y Moo
e
ar re et
rema
e al.
in
h (200
poorl
er 8)
y
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kno
a ed
wn.
Sabi
c
acea
e
e as Pr
o
siste
o
u r to
t
s Prot
e
a eales
a
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d supp
e
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s
ei only
r mod
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le erate
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furth
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v
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e rms
e
th this
r
e posit
c
s ion,
h
a Sabi
t
m acea
.
ee
di will
&
st be
in inclu
J
ct ded
.
iv in a
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)
,
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o
m
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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
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m
m
o
n.

m.
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ns
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(
i
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l
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d
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ic
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er
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rate
8
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5)r for
, this
n mor
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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
t
.

e
( C.
1 N
8 el
2 so
n
2
(2
) 00
, 2)
n
o T
he
m
.limit
cs of
oBux
nacea
se are
.expa
(nded
i. The
nmon
cogelneric
uDidy
dmela
iceae
nhave
gthe
same
disti
D
inctiv
de
ypolle
m
n
eand
lche
amistr
cy as
eat
aleast
epart
of
LBux
eacea
ae,
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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
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ux
g
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ye
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el
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er(
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t c
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(
CORE
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8
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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
8g
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7l
)e
,s
n
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nt
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.&
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r

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r r
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T
e
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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

THE ANGIOSPERM PHYLOGENY GROUP

nom.
dismemberment
of
Celastraceae
cons.
ROSI
needed
to
DSAlthough
maintain
the
Krameriaceae are
Vital
families, of which
monogeneric, they
es
only
are
readily
Jus
Parnassiaceae are
distinguished
well
known,
s.
from
the
would
be
ex
heterogeneous
extensive
and
Ber
Zygophyllaceae;
yield
poorly
cht.
the two are sister
characterized
&
taxa. Combin-ing
families,
and
J.Pr
the two would
Celastraceae s.l.
esl
simply make a
are
better
(18
heterogeneous
characterized than
20)
Zygophyllaceae
Celas-traceae,
Vit
still more so. An
excluding
ace
order is needed
Parnassiaceae (for
for this family
ae
morphology, see
pair as it is placed
Jus
Matthews
&
with
strong
s.
Endress, 2005a).
support as sister to
(17
a clade containing
89)
Oxalidales
more than two
,
Bercht. &
fabid
orders
in
the
no
J.Presl
analysis by Wang
m.
(1820)
et al. (2009).
con
Brunelliace
s.
ae Engl.
Celastrales Link
(1897),
Vitaceae remain(1829)
nom. cons.
isolated
and $Celastraceae
Cephalotac
ordinal status is
R.Br. (1814),
eae
appro-priate. They
nom. cons.
Dumort.
are sister to the
(including
(1829),
fabids + malvids
Lepuropetala
nom. cons.
(rosid I + II) clade
ceae Nakai,
Connaracea
in most recent
Parnassiacea
e R.Br.
analyses,
albeit
e Marti-nov,
(1818),
without
strong
nom. cons.,
nom. cons.
support (reviewed
Pottingeriace
Cunoniacea
in Wang et al.,
ae Takht.)
e R.Br.
Lepidobotryace
2009).
(1814),
ae J.Lonard
nom. cons.
(1950),
nom.
FABI
Elaeocarpa
cons.
DS
ceae Juss.
The limits of
Zygophyllales
ex DC.
Celastraceae are
Link (1829)
(1816),
broadened
$
nom. cons.
because the three
$Krame
*Huaceae
small
families
riaceae
A.Chev.
included
show
Dumort.
(1947)
every sign of
(1829),
making
Oxalidaceae
nom.
Celastraceae
R.Br. (1818),
cons. $
paraphyletic
if
nom. cons.
$Zygop
excluded (Zhang
hyllacea
Huaceae
are
&
Simmons,
tentatively
e R.Br.
2006).
The
included
in
(1814),

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,

Botanical Society, 2009, 161, 105

Journal of 121
the Linnean

APG III 113

n
) that
has
E Chry
disti
u songui
p bala
nace
shin
hr
ae
g
o
woul
featu
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gniti
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eous
on of
a
(see
a
e
Matt
bige
a
hews
neric
n
&
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d
ropla
T Endr
cace
ri ess,
2008
ae is
g
,
for
reaso
o
the
nabl
ni
morp
e.
a
Salic
c holo
acea
e gy of
e are
a this
grou
broa
e
p).
dly
ar
Bhes
draw
e
a
n,
k
i.e.
e (for
inclu
pt merl
y
of
ding
se
Cela
Sam
p
strac
ydac
ar
eae)
eae
at
and
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e
and
asCent
Scyp
, ropl
acus
hoste
al
(for
giace
th
merl
ae
o
y
of
Hutc
u
h.
g Euph
Alth
h orbia
ough
cl ceae)
form
the
e
an
com
ar
isola
bine
ly
ted
d
re
clade
la clade
is
te (Dav
is
et
only
d,
al.,
mod
a
2005
erate
br
;
ly
o
Wur
disti
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nct
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morp
y&
Davi
holo
dr
s,
gical
a
2009
ly
w

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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
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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.
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flora
ceae
and
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nace
ae
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broa
dly
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mite

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ai Med
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ar and
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ct sher
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la s.l.)
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unit.
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c
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oe a
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n

c
l
u
d
i
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g
H
e
t
e
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o
p
y
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i
d
a
c
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a
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E
n
g
l
.
&
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i
l
g
,
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.
c
o
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s
.
,
P
s
i
l
o
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a
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e
a
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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
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g
.
O
B
l
r
i
i
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s
c
)
e V
a oc
e hy
A si
r ac
ea
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e
., A.
n St
o .m Hi
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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

THE ANGIOSPERM PHYLOGENY GROUP

(1858)
least common in
the order. The
,
nom.
chocalycaceae is
sister
taxa
cons.
clear;
Van
Strasburgeri-aceae
Staph
Beusekom-Osinga
and
Ixerbaceae
& van Beusekom yleace
are
two
ae
(1975) included the
monogeneric
last two families in Marti
families that agree
the
first.
All nov
in
several
(1820)
families
are
characters,
,
nom.
morphologically
including
base
similar, although cons.
chro-mosome
they
show
number
and
variation in floral *Strasburgeria
stamen
and
ceae
Soler.
morphology
and
gynoecial
(1908), nom.
embryo sac, in
morphol-ogy etc.;
cons.
particular.
Some
combination is in
(including
combination is in
order. As the order
Ixerbaceae
order,
and
is now defined,
Griseb. ex
Penaeaceae have
Staphyleales
been expanded to
Doweld &
Mart. (1835) is an
include
Reveal)
older name for
Rhynchocalycacea
The addition of
Crossosomatales
e and Oliniaceae;
several families to
Takht. ex Reveal
Penaeaceae s.l. can
Crossosomatales
(1993). It need not
be characterized.
is well justified
be
adopted,
(e.g.
Sosa
&
however.
Crossoso
Chase,
2003;
matales
Soltis
et
al.,
Takht.
2007b; Wang et
ex
al.,
2009),
Picramniales
although
Reveal
Doweld (2001)
monogeneric
(1993)
*Picramniaceae
Guamatelaceae
*Aphlo
Fernando &
are a somewhat
iaceae
Quinn (1995)
surprising
Takht.
Picramniaceae
addition,
(1985)
were
previously
Guamatela
having
Crosso
unplaced rosids,
previously been
somata
but there is now
included
in
ceae
strong support for
Rosaceae (Oh &
Engl.
a position in the
Potter, 2006). For
(1897),
rosid
II/malvid
the most part,
nom.
clade
(Wang
et
relationships
cons.
al.,
2009).
among
families
*Geiss
included
in
olomat
Huerteales
Crossoso-matales
aceae
Doweld (2001)
have not been
A.DC.
*Dipentodonta
suggested before.
(1856)
ceae Merr.
The
order
is
(1941), nom.
*Guam
heterogeneous,
cons.
atelace
although
the
*Gerrardinacea
ae
e Alford (2006)
families are all
S.Oh &
Tapisciaceae
small.
D.Potte
Takht. (1987)
Nevertheless,
r
Matthews
&
(2006)
This
Endress (2005b,
Stachy
assemblage
of
2006) found a
uraceae
three
small
number of floral
J.Agar
families is well
features to be at
dh

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

Botanical Society, 2009, 161, 105

Journal of 121
the Linnean

APG III 115

ugh
ceae

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
H
ei
m
e
rl
(
1
9
3
4
),
n
o
m
.
c
o
n
s.
A
iz
o
a
c
e
a
e
M
a
rt
i
n
o
v
(
1
8
2
0
),
n
o
m
.

co
ns
.
A
m
ar
an
th
ac
ea
e
Ju
ss.
(1
78
9)
,
no
m.
co
ns
.
*
A
na
ca
m
ps
er
ot
ac
ea
e
E
gg
li
&
N
yf
fe
le
r
(2
01
0,
in
p
r
e
s
s
)
A
n
c
i
s
t
r
o

c
l
a
d
a
c
e
a
e
P
l
a
n
c
h
.
e
x
W
a
l
p
.
(
1
8
5
1
)
,
n
o
m
.
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
) .
B
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
)
,
n
o
m
.
c
o
n
s
.

D
i
d
i
e
r
e
a
c
e
a
e
R
a
d
l
k
.
(
1
8
9
6
)
,
n
o
m
.
c

o
n
s
.
D
i
o
n
c
o
p
h
y
l
l
a
c
e
a
e
A
i
r
y
S
h
a
w
(
1
9
5
2
)
,
n
o
m
.
c
o
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r
o
s
e
r
a
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a
l
i
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b.
(1
80
8)
,
no
m.
co
ns
.
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ro
so
ph
yl
la
ce
ae
C
hr
te
k,
Sl
av
k
ov

&
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ud
ni
c
ka
(
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9
8
9
)
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r
a
n
k
e
n
i
a
c
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a
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D
e
s
v
.
(
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1
7
)
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n
o
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.
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o
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s
.
G
i
s
e
k
i
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k
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i
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a
l
o
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.
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o
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a
n
o
(
1
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i
m
e
a
c
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h
i
p
u
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)
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o
p
h
i
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l
d
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8
)

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g
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(
1
8
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5
)
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n
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m
.
c
o
n
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.
*

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n
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a
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R
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f
.
(
1
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)
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e
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m
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.
(
1
8
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)
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.

N
y
c
t
a
g
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n
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c
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J
u
s
s
.
(
1
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)
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(
1
9
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)
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h
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t

o
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.
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)
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.

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c
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s
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.
(
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)
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)
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)

(
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m
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*
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h
.
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r

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m
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nk
(1
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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

THE ANGIOSPERM PHYLOGENY GROUP

Endress
&
cons.
Bittrich,
1993).
Hydrostachy
et al., in press).
Limeaceae
and
aceae Engl.
Expansion
of
Lophiocarpaceae
(1894), nom.
Cactaceae
to
are
segregates
cons.
include all or some
by
Loasaceae
of these smallnecessitated
Juss. (1804),
clades
cannot,recent
nom. cons.
however,
bephylogenetic
justified. Not onlyfindings (Cunoud
Ericales Bercht. &
have the limits ofet al., 2002).
J.Presl (1820)
Cactaceae
beenMolluginaceae s.l.
Actinidiac
stable over theare yet another
eae Engl.
years, but inclusiongroup much in
& Gilg.
of Anacampserosneed of basic
(1824),
and
relativesanatomical,
nom.
(Anacampserotace developmental
cons.
ae)
andand
Balsamina
Portulacaceae s.s. phytochemical
ceae
in
Cactaceaestudy. We still
A.Rich.
(compatible withknow little about
(1824),
of
the best phyloge-relationships
nom.
netic hypotheses)Phytolaccaceae,
cons.
would yield a cladealmost certainly
Clethrace
not
a
characterized
ae
solely
bymonophyletic
Klotzsch
family as curmulticellular
(1851),
axillary
hairs.rently
nom.
Similarly,
inclu-circumscribed.
cons.
its
sion of these twoHowever,
Cyrillacea
groups
anddivision, as in
e Lindl.
Judd
et
al.
(2007),
Talinaceae
in
(1846),
Cactaceae wouldis premature.
nom.
yield
a
clade
cons.
characterized
by
Diapensia
parallelocytic
AST ceae
stomata (probably)
ERID Lindl.
and
fruit
(1836),
S
characters, but the
nom.
latter have beenCornales Link.
cons.
subsequently lost(1829)
Ebenacea
in
Cactaceae Cornaceae
e Grke
(Ogburn
&
Bercht. &
(1891),
Edwards,
2009).
J.Presl
nom.
Anacampserotacea
(1825), nom.
cons.
e has only recently
cons.
Ericaceae
been
described
(including
Juss.
Nyssaceae
(Nyffeler & Eggli,
(1789),
Juss. ex
in
press),
nom.
Dumort.)
recognition
is
cons.
compatible
with Curtisiaceae
Fouquieri
Takht.
(1987)
their phylogenetic
aceae DC.
Grubbiaceae
position.
(1828),
Endl. ex
nom.
Meisn.,
cons.
It has long been (1841), nom.
Lecythida
recognized that the cons.
ceae
limits
of Hydrangeac
A.Rich.
Molluginaceae s.l. eae Dumort.
(1825),
are unclear (e.g. (1829), nom.
nom.

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

are a monogenericalso Mabberley,

5
Samolaceae Raf.2008), although
9
by the break-up ofwe know that this
)
Primulaceae
(ormove will not be
Theophrastaceae univer-sally
I
would have to bewelcomed.
c
extended,
so
a
becoming
less
c
recognizable; see
i
Kllersj,
n
Bergqvist
&
LAMI a
Anderberg, 2000;
IDS c
Sthl & Anderberg,
e
2004, for infor*Boraginaceae
a
mation). The limits
Juss. (1789),
e
of
Myrsinaceae
nom. cons.
were extended, and
(including
those
of
M
Hoplestigmat
Primulaceae
i
aceae Gilg,
correspondingly
e
nom. cons.)
restricted. Given V
r
that the limits of
s
a
the
two
besth
known families in
(
l
the
group,
1
Myrsinaceae and i
8
Primulaceae, have a
5
been substantially c
1
e
changed,
)
apomorphies are a
,
hard to recognize e
(less
so
for
n
Maesaceae
and D
Primulaceae s.s.), a
o
and the group as a n
m
whole
has d
.
numerous
syna- y
pomorphies and is
c
easy to recognize
o
(
so we extend the
n
1
limits
of
s
9
Primulaceae (see
.
Linnean
2009 The Society of
London,

Botanical Society, 2009, 161, 105

Journal of 121
the Linnean

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

Orobanchaceae Vent. (1799), nom. cons.

Paulowniaceae Nakai (1949)
Pedaliaceae R.Br. (1810), nom. cons.
Phrymaceae Schauer (1847), nom. cons.
Plantaginaceae Juss. (1789), nom. cons.
Plocospermataceae Hutch. (1973)
Schlegeliaceae Reveal (1996)
Scrophulariaceae Juss. (1789), nom. cons.
Stilbaceae Kunth (1831), nom. cons.
Tetrachondraceae Wettst. (1924)
*Thomandersiaceae Sreem. (1977)
Verbenaceae J.St.-Hil. (1805), nom. cons.
Note that relationships among many families in
Lamiales, and to a certain extent also their limits,

117

unclear, in the former case despite the sequencing of

the whole plastid genome (Moore et al., in press).
Three families, Icacinaceae, Metteniusaceae and
Oncothecaceae, are to be placed in this general area
of the tree. Furthermore, genera that used to be
included in Icacinaceae s.l. are also to be found here,
although they do not group with Icacinaceae s.s.
(Krehed, 2001); these include Apodytes Arn.,
Cassinopsis Sond. and Emmotum Ham. (= Emmotaceae Tiegh.). All these taxa show similarities to Garryales, and circumscription of that order could easily
be expanded to include them if phylogenetic relationships warranted it. Revised family limits depend on
further phylogenetic work.
Garryales Lindl. (1835)
Eucommiaceae Engl. (1907), nom. cons.
$Garryaceae Lindl. (1834), nom. cons. (including
Aucubaceae Bercht. & J.Presl)
Although Aucubaceae and Garryaceae (both monogeneric) appear distinct, there are several apomorphies for the combined group.
Gentianales Juss. ex Bercht. & J.Presl (1820)
Apocynaceae Juss. (1789), nom. cons.
Gelsemiaceae Struwe & V.A.Albert (1995)
Gentianaceae Juss. (1789), nom. cons.
Loganiaceae R.Br. ex Mart. (1827), nom. cons.
Rubiaceae Juss. (1789), nom. cons.
Lamiales Bromhead (1838)
Acanthaceae Juss. (1789), nom. cons.
Bignoniaceae Juss. (1789), nom. cons.
Byblidaceae Domin (1922), nom. cons.
Calceolariaceae Olmstead (2001)
Carlemanniaceae Airy Shaw (1965)
Gesneriaceae Rich. & Juss. (1816), nom. cons.
Lamiaceae Martinov (1820), nom. cons.
*Linderniaceae Borsch, K.Mll., & Eb.Fisch. (2005)
Lentibulariaceae Rich. (1808), nom. cons.
Martyniaceae Horan. (1847), nom. cons.
Oleaceae Hoffmanns. & Link (1809), nom. cons.

2009 The Linnean Society of

London, Botanical Journal of

the Linnean Society, 2009,
161, 105121

are still unclear. Some of us would prefer a vastly

expanded circumscription of Scrophulariaceae, far
beyond what it has ever included, whereas others
are not so inclined. The limits of Plantaginaceae
have been further restricted since APG II by the
recognition of the family of small herbs with rather
distinctive stem anatomy and floral morphology
(e.g. Linderniaceae), and Thomandersia has been
removed from Acanthaceae as the monogeneric
Thomandersiaceae (Wortley, Harris & Scotland,
2007).
Solanales Juss. ex Bercht. & J.Presl (1820)
Convolvulaceae Juss. (1789), nom. cons.
Hydroleaceae R.Br. ex Edwards (1821)
Montiniaceae Nakai (1943), nom. cons.
Solanaceae Juss. (1789), nom. cons.
Sphenocleaceae T.Baskerv. (1839), nom. cons.

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

THE ANGIOSPERM PHYLOGENY GROUP

include the monopress). It is likely
generic
to be sister to
P
Donatiaceae
is
Asterales or, more
h
supported
by
probably, to all
e
morphology and
campanulids apart
l
geography,
and
from
Asterales
l
the
expanded
and Aquifoliales.
i
Campanulaceae
Even if sister to
n
have
strong
Asterales,
a
support
in
inclusion in that
c
molecular
studies
order would make
e
and
are
well
the
latter
a
characterized
distinctly
more
e
morphologically.
heterogeneous;
Relationships
separate ordinal
T
within
status is needed.
a
Campanulaceae
k
s.l.
are
still
h
Bruniales
unclear (Tank &
t
Dumort. (1829)
Donoghue,
in
.
Bruniaceae
press), and a
R.Br. ex DC.
future attempt to
(
(1825), nom.
recog-nize
1
cons.
Lobeliaceae might
9
Columelliacea
either result in a
6
e D.Don
clade
poorly
7
(1828), nom.
supported
)
cons. (includmorphologically
ing
or
entail
the
R
Desfontainia
recognition of yet
o
ceae Endl.,
other families in
u
nom. cons.)
this complex.
s
An order is
s
needed
for the
e
Escalloniales
two
families
a
R.Br. (1835)
above.
c
Escalloniacea
Winkworth,
e
e R.Br. ex
Lundberg
&
a
Dumort.
Donoghue
(2008)
e
(1829), nom.
found
some
cons.
support for a
D
(including
position sister to
Eremosynac
C
Asterales
and
eae Dandy,
.
Polyosmacea
Tank & Donoghue
e
(in press) found
(
Blume,
stronger support
1
Tribelaceae
for a position
8
Airy Shaw)
sister
to
the
3
Paracryphiales
9
This
is
a
)
Dipsacales
heterogeneous
$Stylidiaceae group of genera
Apiales
clade;
ordinal status is
R.Br. (1810), that forms a wellappropriate.
nom. cons. supported clade,
Columelliaceae
(including
but
one
of
are
broadly
Donatiaceae uncertain position
circumscribed
B.Chandler, and within which
because
nom. cons.) relationships are
Desfontainiaceae
poorly supported
Expansion
of(Tank
have much in
&
Stylidiaceae
toDonoghue,
common
with
in

them; both familiesDipsacales Juss.

are Andean and ex Bercht. &
J.Presl (1820)
monogeneric.
Adoxaceae
E.Mey. (1839),
Paracryphiales
nom. cons.
Takht. ex Reveal
Caprifoliaceae
(1992)
Juss. (1789),
Paracryphiacea
nom. cons.
e Airy Shaw
[including
(1965)
Diervillaceae
(including
Pyck,
*Quintiniacea
Dipsacaceae
e Doweld,
Juss., nom.
Sphenostemo
cons.,
naceae
Linnaeaceae
P.Royen &
Backlund,
Airy Shaw
Morinaceae
(1972))
Raf.,
Valerianacea
Although these
e
Batsch,
three families are
nom. cons.]
at
first
sight
strikingly different, A
broad
they have severalcircumscription of
characters
inCaprifoliaceae is
common and formadopted here as it
a
stronglyis
widely
supported
clade
preferred (Judd et
(Tank
&
al.,
2007;
Donoghue,
in
Mabberley, 2008).
press); all are
The
expanded
monogeneric and
family is well
from
the
characterized, but
southwestern
half the clades it
Pacific.
includes
are
Combination is in
poorly
order (see also
characterized
Myrtales,
morphologically.
Crossosomatales).
Tank & Donoghue
(in press) foundApiales Nakai
100%
bootstrap(1930)
support
for
a A
position
of p
Paracryphiales as i
sister
to a
c
Dipsacales.
e
a
e
L
i
n
d
l
.
(
1
8

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
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of
ea
members included
e
formerly
in
R.
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81
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OF
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UNCER
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TAIN
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ON
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Hu
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e Takhtajan
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Philipson &Cynomoriacea
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Melanophylla Lindl. (1833),
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Airy Shaw) Gumillea Ruiz
& Pav.
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.
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ter
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Ch
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