Botanical Journal of the Linnean Society, 2016, 181, 1–20.

With 1 figure

An update of the Angiosperm Phylogeny Group

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classification for the orders and families of flowering
plants: APG IV
THE ANGIOSPERM PHYLOGENY GROUP1*
1
Recommended citation: APG IV (2016). This paper was compiled by James W. Byng, Mark W. Chase, Maarten J. M.
Christenhusz, Michael F. Fay, Walter S. Judd, David J. Mabberley, Alexander N. Sennikov, Douglas E. Soltis, Pamela S. Soltis
and Peter F. Stevens, who were equally responsible and listed here in alphabetical order only, with contributions from Barbara
Briggs, Samuel Brockington, Alain Chautems, John C. Clark, John Conran, Elspeth Haston, Michael M€ oller, Michael Moore,
Richard Olmstead, Mathieu Perret, Laurence Skog, James Smith, David Tank, Maria Vorontsova and Anton Weber.
Addresses: M. W. Chase, M. J. M. Christenhusz, M. F. Fay, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, UK;
J. W. Byng, M. J. M. Christenhusz, Plant Gateway, 5 Talbot Street, Hertford, Hertfordshire SG13 7BX, UK; J. W. Byng, School
of Biological Sciences, University of Aberdeen, Aberdeen AB24 3UU, UK; M. W. Chase, University of Western Australia, 35
Stirling Highway, Crawley, Western Australia 6009, Australia; W. S. Judd, D. E. Soltis, Department of Biology, University of
Florida, Gainesville, FL 32611-8525, USA; D. J. Mabberley, Wadham College, University of Oxford, UK; Universiteit Leiden and
Naturalis Biodiversity Center, Leiden, the Netherlands; Macquarie University and National Herbarium of New South Wales,
Sydney, Australia; A. N. Sennikov, Botanical Museum, Finnish Museum of Natural History, PO Box 7, FI-00014, Helsinki,
Finland and Komarov Botanical Institute, Prof. Popov 2, RU-197376, St. Petersburg, Russia; D. E. Soltis, P. S. Soltis, Florida
Museum of Natural History, University of Florida, Gainesville, FL 32611-7800, USA; P. F. Stevens, Department of Biology,
University of Missouri-St. Louis and Missouri Botanical Garden, PO Box 299, St. Louis, MO 63166-0299, USA.

Received 10 January 2016; revised 17 January 2016; accepted for publication 17 January 2016

An update of the Angiosperm Phylogeny Group (APG) classification of the orders and families of angiosperms is
presented. Several new orders are recognized: Boraginales, Dilleniales, Icacinales, Metteniusiales and Vahliales.
This brings the total number of orders and families recognized in the APG system to 64 and 416, respectively. We
propose two additional informal major clades, superrosids and superasterids, that each comprise the additional
orders that are included in the larger clades dominated by the rosids and asterids. Families that made up
potentially monofamilial orders, Dasypogonaceae and Sabiaceae, are instead referred to Arecales and Proteales,
respectively. Two parasitic families formerly of uncertain positions are now placed: Cynomoriaceae in
Saxifragales and Apodanthaceae in Cucurbitales. Although there is evidence that some families recognized in
APG III are not monophyletic, we make no changes in Dioscoreales and Santalales relative to APG III and leave
some genera in Lamiales unplaced (e.g. Peltanthera). These changes in familial circumscription and recognition
have all resulted from new results published since APG III, except for some changes simply due to nomenclatural
issues, which include substituting Asphodelaceae for Xanthorrhoeaceae (Asparagales) and Francoaceae for
Melianthaceae (Geraniales); however, in Francoaceae we also include Bersamaceae, Ledocarpaceae,
Rhynchothecaceae and Vivianiaceae. Other changes to family limits are not drastic or numerous and are mostly
focused on some members of the lamiids, especially the former Icacinaceae that have long been problematic with
several genera moved to the formerly monogeneric Metteniusaceae, but minor changes in circumscription include
Aristolochiaceae (now including Lactoridaceae and Hydnoraceae; Aristolochiales), Maundiaceae (removed from
Juncaginaceae; Alismatales), Restionaceae (now re-including Anarthriaceae and Centrolepidaceae; Poales),
Buxaceae (now including Haptanthaceae; Buxales), Peraceae (split from Euphorbiaceae; Malpighiales),
recognition of Petenaeaceae (Huerteales), Kewaceae, Limeaceae, Macarthuriaceae and Microteaceae (all
Caryophyllales), Petiveriaceae split from Phytolaccaceae (Caryophyllales), changes to the generic composition of
Ixonanthaceae and Irvingiaceae (with transfer of Allantospermum from the former to the latter; Malpighiales),
transfer of Pakaraimaea (formerly Dipterocarpaceae) to Cistaceae (Malvales), transfer of Borthwickia,
Forchhammeria, Stixis and Tirania (formerly all Capparaceae) to Resedaceae (Brassicales), Nyssaceae split from
Cornaceae (Cornales), Pteleocarpa moved to Gelsemiaceae (Gentianales), changes to the generic composition of
Gesneriaceae (Sanango moved from Loganiaceae) and Orobanchaceae (now including Lindenbergiaceae and

*E-mail: m.chase@kew.org

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20 1
2 THE ANGIOSPERM PHYLOGENY GROUP

Rehmanniaceae) and recognition of Mazaceae distinct from Phrymaceae (all Lamiales). © 2016 The Linnean
Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20

ADDITIONAL KEYWORDS: Apodanthaceae – Aristolochiaceae – Boraginales – Cistaceae –

Cynomoriaceae – Dasypogonaceae – Dilleniales – Francoaceae – Gesneriaceae – Icacinales –

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Metteniusales – Orobanchaceae – Phrymaceae – Phytolaccaceae – Resedaceae – Restionaceae –
Sabiaceae – Santalales – Vahliales.

sense might be problematic, such as Dioscoreaceae

HISTORICAL INTRODUCTION
(Caddick et al., 2002), Flacourtiaceae (Chase et al.,
In 1998, the first Angiosperm Phylogeny Group (APG) 2002), Lamiaceae/Verbenaceae (Wagstaff & Olm-
classification of the orders and families of flowering stead, 1997), Loganiaceae (Backlund, Oxelman &
plants (which we will term APG I; APG, 1998) was Bremer, 2000), Malvaceae (Judd & Manchester, 1997;
published, and this classification initiated a new Bayer et al., 1999), Rutaceae (Chase, Morton & Kal-
approach to this long tradition. APG I was not written lunki, 1999) and others. These early studies of puta-
by one or two authoritative individuals; rather the tively problematic families resulted in mostly clear-
APG process tried to produce a consensus classifica- cut solutions, especially if one followed the principles
tion that reflected results and opinions of experts in of Backlund & Bremer (1998; Vences et al., 2013 pre-
many groups of flowering plants. The initial focus sented a zoological perspective on these same issues).
was to produce a classification of families in orders, Again, gaining a consensus was relatively straightfor-
without too much emphasis on the issue of family ward, and APG II (2003) tried to make this easier by
delimitation; in 1998, few families had been appropri- offering APG users optional circumscriptions, nar-
ately studied, and so such issues had limited consider- rower and broader (a ‘bracketed’ system), permitting
ation in APG I. Exceptions were families, such as any permutation thereof and still allowing authors to
Saxifragacaeae (Morgan & Soltis, 1993), Geraniaceae claim that they were ‘following APG’. Delimitation of
(Price & Palmer, 1993), Liliaceae (Chase et al., 1995), families was clearly becoming a major issue, and the
Onagraceae (Conti, Fischbach & Sytsma, 1993) and use of the bracketed system was, in addition to an
Ericaceae (Judd & Kron, 1993; Kron & Chase, 1993), attempt to maintain a broad consensus of support,
that had been the focus of early molecular studies, focused much more on family delimitation and the
some of them due to their suspected polyphyletic nat- issue of lumping versus splitting.
ure (e.g. Saxifragaceae sensu Cronquist, 1981). In response to negative reactions received by the
Because the rule of priority does not apply at the level compilers on the use of the bracketed system, this
of order, the biggest issue in APG I was standardiza- usage was abandoned in APG III (2009). In most,
tion of names being applied to orders so that research- but not all, cases the broader circumscriptions
ers (many of them using molecular techniques) implied by the bracketing were accepted. As
studying similar sets of families were not using differ- reviewed by Wearn et al. (2013) and Christenhusz
ent names. Prevention of chaos was the objective, and et al. (2015), this lumping approach was made in an
consensus was relatively easily reached. The sum- effort to simplify the parts of a classification that
mary consensus tree (fig. 1, p. 535) provided in APG I users emphasize, principally orders and families, an
was highly unresolved, an indication of the prelimi- approach that has generally received support. An
nary nature of what was known at that time about additional effort to assess support from both tax-
higher-level (interordinal) relationships, even though onomists and users of classification for broader ver-
the composition of what were then considered orders sus narrower circumscriptions was made by
was reasonably clear. conducting an online survey in August 2014 (Chris-
The general scheme of the arrangement of major tenhusz et al., 2015), with the realization that any
groups was also clear: a grade of isolated taxa (the survey may have biases due to the way questions are
ANA grade, or ANITA grade as it was then called) phrased.
leading to the major radiation of angiosperms, a
clade of all monocots, a clade of magnoliid families
and a large eudicot (tricolpate) clade composed of
REVIEW
several small clades and two major groups, rosids
and asterids, each composed of two major subclades. Higher-level classification of angiosperms has
As the general framework of angiosperm relation- received continuing attention since APG III (2009)
ships became clearer, the focus started to shift and enough progress has been made that an update
toward issues of family delimitation, with an empha- to the APG classification is warranted. Several
sis on those that most angiosperm taxonomists had a important studies have been published since 2009

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
 APG IV 3

Amborellales
Nymphaeales
Austrobaileyales

Angiosperms
Magnoliales
Laurales Magnoliids
Piperales
Canellales

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Chloranthales
Arecales
Poales Commelinids
Commelinales
Zing iberales

Monocots
Asparagales
Liliales
Diosco reales
Pandanales
Petros aviales
Alis matales
Acorales
Ceratophyllales
Ranunculales
Proteales
Trochodendrales

Eudicots
Buxales
Gunnerales
Fabales
Rosales
Fagales
Cucurbitales
Oxalidales Fabids
Malpighiales
Celastrales
Superrosids Zygophyllales
Geraniales
Rosids Myrtales
Crossosomatales
Picramniales
Malvales Malvids
Brassicales
Huerteales
Sapindales
Vitales
Saxifragales
Figure 1. Interrelationships of the †Dilleniales
APG IV orders and some families Berb eridopsidales
Santalales
supported by jackknife/bootstrap Caryophyllales
Superasterids

percentages >50 or Bayesian Cornales

Ericales
posterior probabilities >0.95 in Aquifoliales
large-scale analyses of angiosperms. Asterales
Asterids

Escalloniales
See text for literature supporting Bruniales Campanulids
these relationships. The alternative Apiales
Dipsacales
placements representing Paracryphiales
incongruence between nuclear/ Solanales
Lamiales
mitochondrial and plastid results †Vahliales
for the Celastrales/Oxalidales/ Gentianales Lamiids
Malpighiales (COM) clade are †Boraginales
Garryales
indicated by slash marks (\\). †Mettenius ales
†Orders newly recognized in APG. †Icacinales

(APG III), particularly those of Soltis et al. (2011), Researchers have speculated about what analyses
Ruhfel et al. (2014) and Stull et al. (2015). Soltis of low-copy nuclear genes would reveal about plant
et al. (2011) used 17 genes from all three genomes relationships and whether these relationships would
for 640 angiosperm taxa, whereas Ruhfel et al. be different from those portrayed so far by plastid,
(2014) used 78 protein-coding plastid genes for 360 mitochondrial and nuclear ribosomal genes. Nuclear
green plant taxa (including green algae). Both analy- data, particularly low-copy genes, have so far been
ses reached similar general conclusions for the poorly represented in broader phylogenetic studies of
angiosperms. Stull et al. (2015) concentrated on the the angiosperms. Morton (2011) surveyed xanthine
lamiids, but this was the clade in which the greatest dehydrogenase (Xdh) for 247 genera of seed plants
uncertainty existed, particularly with the former and obtained results generally congruent with those
Icacinaceae, which had been known to be poly- of previous studies, although the branching order
phyletic (Savolainen et al., 2000). within some larger clades was different from other

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
4 THE ANGIOSPERM PHYLOGENY GROUP

studies. Zeng et al. (2014) and Wickett et al. (2014) & Brummitt, 2003) and some authors strictly prefer
both analysed low-copy nuclear genes (59 and 852 genes, the traditional versions (e.g. Compositae vs. Aster-
respectively), but relatively few angiosperms (60 and 37, aceae). At the Royal Botanic Gardens, Kew, for
respectively, the latter focused on all green plants), example, Compositae and Leguminosae are formally
and reached similar conclusions about relationships to endorsed, whereas the reverse is true for Apiaceae

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those found in the majority of earlier studies. (not Umbelliferae), Arecaceae (not Palmae), Brassi-
Although the results using low-copy nuclear genes caceae (not Cruciferae), Clusiaceae (not Guttiferae),
may not substantially alter our ideas of the major Lamiaceae (not Labiatae) and Poaceae (not Grami-
framework of relationships within the angiosperms, neae). These alternative names are considered as not
there is at least one consistent and significant differ- based on a generic name, and they are attached to
ence. Celastrales, Oxalidales and Malpighiales (the particular genera by the means of a special provision
COM clade), in general found in the fabid clade of in the International Code of Nomenclature for algae,
rosids (rosid I) based on the mostly plastid DNA fungi and plants (McNeill et al., 2012; Art. 18.5). We
results published up to 2011, are instead members of list these alternative names here (in parentheses) for
the malvid clade (rosid II) in trees inferred from low- the first time because they are of equal status in the
copy nuclear and mitochondrial genes. This is consis- Code, continue to receive wide use in the literature
tent with the nuclear results of Morton (2011), Zeng and are preferred by many working on the groups
et al. (2014) and Wickett et al. (2014) and mitochon- concerned; see Mabberley (2008: xi–xii) for further
drial results of Zhu et al. (2007) and Qiu et al. (2010). discussion.
Sun et al. (2015) reviewed the history of these incon- The suprageneric names appearing in Martinov
gruent results and added additional studies of mito- (1820) have been subject to extensive debate, and a
chondrial and nuclear genes. It is possible that some proposal to treat all names as not validly published
sort of horizontal transfer of plastid DNA, perhaps in that book has been made recently (Sennikov et al.,
via ancient hybridization, produced this incongruence 2015). If this proposal is accepted, the authorship
(Sun et al., 2015). We have indicated this incongru- and dates of such names will be changed, affecting
ence in Figure 1. It is not yet clear if this incongru- at least the name Acoraceae (all other Martinov
ence extends to Zygophyllaceae, which fell as sister to names are conserved, and their place of publication
the rest of the fabid clade (including the COM clade) can only be changed by the means of proposals to
in plastid analyses in Sun et al. (2015). amend entries of conserved names; Art. 14.15). Addi-
In this update of APG, there are some changes from tionally, several familial names are credited to Van
APG III as a result of placements of some genera that Tieghem, although they appeared not in Van Tie-
required erection of new families, and we recognize ghem’s work but in reviews of his articles published
several new orders as a result of studies incorporating in Just’s Botanischer Jahresbericht. Because of con-
many genes/whole plastid genomes (Soltis et al., troversies connected to acceptance and authorship of
2011; Ruhfel et al., 2014; Stull et al., 2015), for exam- such publications, they have also been proposed to be
ple Boraginales, Dilleniales, Icacinales and Mette- treated as inappropriate for valid publication (Sen-
niusales (see below). We deviate here from previous nikov et al., 2015). If this proposal is accepted, the
APG papers in placing the families in the linear order relevant familial names should be credited to later
of Haston et al. (2009; LAPG) and provide comments authors who accepted Van Tieghem’s names and ful-
on changes and other issues in the text below, thus filled the conditions for their valid publication.
keeping the linear sequence of orders and families Two entries of conserved familial names, i.e. Actini-
intact. For a formal, higher-level classification of diaceae and Eucommiaceae, were found (Reveal,
plants, see Cantino et al. (2007) and Chase & Reveal 2010) to have been published earlier than recorded in
(2009), which can still be applied to this version of the list of conserved names. The name Actinidiaceae
APG. Recently, linear orders and revised classifica- was also published with a different authorship
tions have been published for ferns and lycopods (Actinidiaceae Engl. & Gilg, not Gilg & Werderm. as
(Smith et al., 2006; Christenhusz & Chase, 2014) and in Wiersema et al., 2015). More additions affecting
gymnosperms (Christenhusz et al., 2011), which pro- conserved familial names are from Batsch (1794),
vide companion classifications for the remainder of which is to be considered as the place for valid publi-
the vascular plant flora. cation of Melanthiaceae and Primulaceae, both
accepted and conserved with the authorship of
‘Batsch ex Borkh. 1797’ but validly published in 1794
by a reference in the introduction of that book to the
A NOTE ON FAMILY NAMES
corresponding descriptions in Batsch (1786). These
Alternative names for eight flowering plant families entries can be corrected by means of special proposals
have been extensively discussed (reviewed by McNeil to avoid current discrepancies in the databases.

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
 APG IV 5

Another issue is standardization of the authorship MONOCOTS

of conserved familial names, which is regulated by
Since APG III (2009), little alteration of our under-
Art. 46 but not covered by Art. 14.15. At present,
standing has been achieved among monocots, but
entries of conserved familial names have inconsistent
several relatively minor changes are proposed here.
statements of authorship when a taxon was described

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First, we place Dasypogonaceae in Arecales on the
by one author but that name was validly published
basis of Barrett et al. (2016), in which they received
later by another author. In the list of conserved famil-
moderate to high support as sister to Arecaceae. Pre-
ial names, such names are attributed either to original
vious studies with much sparser taxonomic sampling
authors (e.g. Theaceae Mirb.), presumed validating
did not strongly support this relationship (Givnish
authors (e.g. Asteraceae Martinov) or both authors
et al., 2010; Ruhfel et al., 2014), even though they
connected with ‘ex’ (e.g. Ancistrocladaceae Planch. ex
placed Dasypogonaceae as sister to Arecaceae.
Walp.). We follow the authorship as attributed by
In Alismatales, we recognize here Maundiaceae
Wiersema et al. (2015), in anticipation that this will
because the single genus, Maundia F.Muell., has a
be standardized in the next edition of the International
non-exclusive relationship with Juncaginaceae (Von
Code of Nomenclature for algae, fungi and plants.
Mering & Kadereit, 2010; Les & Tippery, 2013), in
which it was previously placed (APG III, 2009). Erec-
tion of another monogeneric family in this order in
ANGIOSPERM CLASSIFICATION: AN which the alismatid families (not including Araceae)
UPDATE are already numerous and small might seem unwar-
ranted, but the online survey (Christenhusz et al.,
By way of general comment on our philosophy of
2015) found little support for the alternative, namely
adopting changes to the APG classification, we have
expansion of Juncaginaceae to include Potamoget-
followed here a conservative approach of accepting
onaceae, Zosteraceae, Cymodoceacee, Ruppiaceae,
only changes due to new phylogenetic studies. With-
Posidoniaceae and Maundiaceae. The simplest solu-
out new results demonstrating a well-supported need
tion to the problem posed by Maundia is the addition
for change, we have maintained the APG III classifi-
of another family to Alismatales.
cation. There are at least two cases in which the
In Dioscoreales, we maintain the circumscription
APG IV classification does not reflect the results of
of the families provided in APG III (2009), but we
published studies, Dioscoreales and Santalales, and
admit that several studies (Merckx et al., 2009; Mer-
in these cases there are either conflicting results
ckx, Huysmans & Smets, 2010; Merckx & Smets,
among the published studies or insufficient support
2014) have indicated that Thismia Griff. and its rela-
for evaluating what possible altered familial circum-
tives and Burmannia L. and related genera do not
scriptions might be possible or preferable, respec-
form a clade. Those authors recommended that This-
tively. In these cases, we await future resolution
miaceae, Burmanniaceae and Taccaceae be rein-
before altering APG III.
stated to reflect their estimates of relationships for
We place Chloranthales on a polytomy with the mag-
these taxa. Caddick et al. (2002), upon which the
noliid and eudicots/monocots/Ceratophyllaceae clades
APG III circumscriptions were based, had earlier
because several recent studies (e.g. Wickett et al.,
found good support for the relationships as recog-
2014; Zeng et al., 2014) have not placed them with the
nized in APG (2003, 2009), and Hertweck et al.
magnoliids, as was indicated in APG III (2009). Sup-
(2015) reaffirmed this relationship in their analysis.
port for Chloranthales as sister to the magnoliids was
We hope that future studies will resolve the incon-
also low (bootstrap support 61–69%) in Ruhfel et al.
gruence reported in the literature for this order, and
(2014).
we will make any necessary changes to familial
There are no alterations among the ANA grade or
circumscription at that time.
the magnoliid families and orders, except for inclu-
To make the name Asphodelaceae available for use
sion of Hydnoraceae and Lactoridaceae in Aris-
when this family in the strict sense is combined with
tolochiaceae due to paraphyly of the last (Massoni,
Xanthorrhoeaceae, conservation of Asphodelaceae
Forest & Sauquet, 2014). It has been known that
was proposed (Klopper, Smith & van Wyk, 2013) and
Lactoris Phil. was embedded in Aristolochiaceae (Qiu
approved by the Nomenclature Committee for Vascu-
et al., 2005; Wanke et al., 2007), but this placement
lar Plants (Applequist, 2014). This action will restore
was considered by some to be an artefact due to a
the priority of Asphodelaceae over Xanthorrhoeaceae
long-branch problem. No study has yet supported
as soon as conservation is approved by the General
this hypothesis, so it seems appropriate to make this
Committee and then the Nomenclature Section of
change in circumscription. Hydnoraceae have also
the XIX International Botanical Congress in Shen-
recently been shown to be nested in Aristolochiaceae
zhen, 2017.
(Naumann et al., 2013; Massoni et al., 2014).

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
6 THE ANGIOSPERM PHYLOGENY GROUP

In Poales, there have been conflicting estimates of gales as sister to the rosid clade, and this more inclu-
relationships among Anarthriaceae, Centrolepi- sive clade, i.e. Saxifragales + rosids, is here referred to
daceae and Restionaceae (reviewed by Briggs, as the superrosids (following Soltis et al., 2011).
Marchant & Perkins, 2014). To stabilize the taxon-
omy of this order, we enlarge Restionaceae to

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re-include Anarthriaceae and Centrolepidaceae so
ROSIDS
that, regardless of the outcomes of future studies,
the family name will remain the same. Few changes to family circumscription have been
made among rosids relative to APG III. The endopar-
asitic Apodanthaceae are now placed in Cucurbitales
(Filipowicz & Renner, 2010). The sequence of families
EUDICOTS
in Malpighiales diverts from that of LAPG (Haston
The classification of Ranunculales and Trochoden- et al., 2009), because we now have a much better
drales remains the same as in APG III (2009). We understanding of interfamilial relationships in that
move Sabiaceae into Proteales on the basis of strong order (Soltis et al., 2011; Xi et al., 2012; Endress,
support found by Sun et al. (2016). Bootstrap support Davis & Matthews, 2013). Also in Malpighiales, there
for this placement was not strong in earlier studies is one newly recognized family, Peraceae, the poten-
(Ruhfel et al., 2014, 63%; Soltis et al., 2011, 59%). In tial need for which was discussed in APG III (2009),
Buxales, we broaden the limits of Buxaceae to due to the position of Rafflesiaceae as sister to the
include Haptanthaceae (Buxaceae already included rest of Euphorbiaceae, minus Pera Mutis and rela-
Didymelaceae in APG III, 2009). Shipunov & Shipu- tives (Davis et al., 2007). This family is now accepted
nova (2011) found that Haptanthus Goldberg & here as Peraceae (Endress et al., 2013). In addition,
C.Nelson was embedded in Buxaceae, possibly sister two changes to familial circumscription are needed.
to Buxus L., so its inclusion in that family is Allantospermum Forman has historically alternated
indicated. between Ixonanthaceae and Irvingiaceae, but most
recently has been considered in Ixonanthaceae (Byng,
2014; Kubitzki, 2014). Recently, J.W. Byng (unpubl.
data) has shown Allantospermum to be sister to the
CORE EUDICOTS (NEITHER ROSIDS NOR
rest of Irvingiaceae rather than Ixonanthaceae.
ASTERIDS)
In Huerteales, Petenaeaceae (Christenhusz et al.,
In this set of clades, only two changes are made, 2010) are added as a new family. In Geraniales,
neither affecting familial circumscriptions. On the Francoaceae must be substituted for Melianthaceae,
basis of results in Soltis et al. (2011) and Ruhfel due to nomenclatural priority, and we include Vivi-
et al. (2014), recognition of monofamilial Dilleniales aniaceae in Francoaceae on the basis of Sytsma,
is warranted. However, in the former they are well Spalink & Berger (2014). Exact relationships among
supported as sister to the large superasterid clade, Francoaceae s.s. (Francoa Cav., Greyia Hook. &
whereas in the latter they are well supported as sis- Harv. and Tetilla DC.), Melianthaceae (Bersama
ter to the large superrosid clade. Due to this conflict, Fresen. and Melianthus L.) and Ledocarpaceae (for
here we do not include them in either larger clade which Vivianiaceae is a later synonym, contrary to
(Fig. 1). In the linear order presented here, the posi- its use in APG III; Balbisia Cav., Rhynchotheca Ruiz
tion of Dilleniales does not exactly accord with their & Pav., Viviania Cav. and Wendtia Meyen) are
phylogenetic position among the eudicots, but this uncertain, with contradictory relationships in recent
set of core eudicots is paraphyletic to rosids plus papers (Palazzesi et al., 2012; Sytsma et al., 2014).
asterids, thus making the sequence of the linear We opt to stabilize APG by recognizing the broader
order arbitrary as long as they are excluded from circumscription so that no matter which relationship
these two larger groups. proves to be the most robust the family name recog-
The other change is the position of Cynomoriaceae, nized does not change.
for which the evidence has been weak and contradic- Alteration of family limits for Sapindaceae (Sapin-
tory in published studies (reviewed in APG III, 2009; dales) was proposed by Buerki et al. (2010) to preserve
and Qiu et al., 2010). Recently, S. Bellot & S. Renner the long-recognized temperate families, Aceraceae and
(unpubl. data) showed that Cynomoriaceae are well Hippocastanaceae. To accomplish this required recog-
supported as members of Saxifragales, although their nition of a new family, Xanthocerataceae, which
exact position in that order is not yet clear. Vitales, on Buerki et al. (2010) published as Xanthoceraceae.
the basis of Soltis et al. (2011) and Ruhfel et al. Relationships in Sapindaceae have been known since
(2014), are again considered in the rosid clade. Both of Harrington et al. (2005) and, given our statement of
these analyses also supported the position of Saxifra- philosophy (above), we do not alter circumscription of

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
 APG IV 7

Sapindaceae because no new phylogenetic information maceae (Feodorova et al., 2010; Patchell, Roalson &
has become available that addresses this issue of Hall, 2014), leaving perhaps only a single genus in
altered family limits. Contrary to the viewpoint of that family. Two genera of Capparaceae (Keithia
Buerki et al. (2010), Sapindaceae s.l. are easily diag- Spreng. and Poilanedora Gagnep.) are a poor mor-
nosed morphologically (Judd et al., 2016). phological fit with their pentamerous flowers. They

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In Malvales, circumscription of Cistaceae, Diptero- most certainly belong elsewhere, and we include
carpaceae and Sarcolaenaceae requires a minor them at the end as unplaced genera.
adjustment, but the potential for further change
should be noted for this set of families. In an analysis
based only on rbcL (Ducousso et al., 2004), Pakarai-
SUPERASTERIDS
maea Maguire & P.S.Ashton (placed in its own sub-
family of Dipterocarpaceae) is sister (88% bootstrap In Santalales, we confront a difficult question about
support) to Cistaceae, and Monotes A.DC. and Pseu- how best to represent the phylogenetic results
domonotes Maguire & P.S.Ashton (subfamily Mono- obtained by Mal
ecot & Nickrent (2008) and Der &
toideae of Dipterocarpaceae) are weakly supported Nickrent (2008), as summarized in Nickrent et al.
(62%) as sister to Sarcolaena Thouars plus Leptolaena (2010), Su et al. (2015) and J.W. Byng (unpubl. data).
Thouars (Sarcolaenaceae; 97%) and Dipterocar- APG III (2009) reported the results of the two phyloge-
poideae (84%). Here, we propose to include Pakarai- netic papers (Der & Nickrent, 2008; Mal
ecot & Nick-
maea in an expanded Cistaceae. Sarcolaenaceae rent, 2008; as summarized in Nickrent et al., 2010),
might also need to be included in Dipterocarpaceae; but refrained from making any changes to the classifi-
they share many morphological, anatomical and cation. Mal
ecot & Nickrent (2008; as summarized in
chemical characters and in Ducousso et al. (2004) are Nickrent et al., 2010) split ‘Olacaceae’ into eight
sister to Dipterocarpoideae to the exclusion of Mono- families: Aptandraceae, Coulaceae, Erythropalaceae,
toideae of Dipterocarpaceae. We refrain from making Octoknemaceae, Olacaceae s.s., Schoepfiaceae, Strom-
further changes in this group of families until a more bosiaceae and Ximeniaceae. Additionally, Der &
comprehensive study (in terms of data and taxa) has Nickrent (2008; as summarized in Nickrent et al.,
been concluded. Perhaps it would be better to combine 2010) proposed recognition of seven families in the
all of these into a single family, given that the limits group recognized as Santalaceae in APG III (2009):
of neither Cistaceae nor Dipterocarpaceae would be Amphorogynaceae, Cervantesiaceae, Comandraceae,
consistent with past circumscriptions. The continued Nanodeaceae, Santalaceae s.s., Thesiaceae and Vis-
use of Dipterocarpaceae (currently used for the eco- caceae. However, strong support for these relation-
nomically most significant group) could be achieved ships is lacking, particularly in ‘Olacaceae’. We
by superconservation of the name Dipterocarpaceae, therefore here opt to maintain the APG III (2009) sta-
as Cistaceae currently has nomenclatural priority and tus quo in Santalales until additional data can be
is a conserved name. brought to bear on this problematic clade. Further-
In Brassicales, the generic composition of Cleo- more, a recent study (J.W. Byng, unpubl. data) places
maceae and Capparaceae has continued to be dimin- Balanophoraceae s.l. as a monophyletic group in ‘San-
ished by studies finding that the genera belong talaceae’, in contrast to Su et al. (2015), where Balano-
elsewhere, with Koeberlinia Zucc. (Koeberliniaceae), phoraceae were divided into two clades. We
Pentadiplandra Baill. (Pentadiplandracee) and acknowledge that our use of ‘Olacaceae’ and ‘Santala-
Setchellanthus Brandegee (Setchellanthaceae) hav- ceae’ does not refer to monophyletic groups and thus
ing already been placed in their own families in APG maintain the families as they were in APG III, but in
III (2009). Su et al. (2012) showed that Borthwickia the linear sequence we move Balanophoraceae next to
W.W.Sm., Forchhammeria Liebm., Stixis Lour. and ‘Santalaceae’, in which they appear to be embedded.
Tirania Pierre are collectively paraphyletic to Rese- Familial delimitation in Caryophyllales continues
daceae and described Borthwickiaceae, whereas a to generate taxonomic conundrums focused on three
separate Stixidaceae (as ‘Stixaceae’) had been previ- problematic sets of families, although the nature of
ously proposed by Doweld & Reveal (2008). Here we these problems is different in each case (reviewed by
include Borthwickiaceae and Stixidaceae in an Hern
andez-Ledesma et al., 2015). The first centres
expanded Resedaceae, members of which share some on Phytolaccaceae and their relationship to Nyctagi-
morphological characters (e.g. flowers with many sta- naceae, which has long posed problems. Genera pre-
mens), although some share more characters with viously associated with Phytolaccaceae but now with
Gyrostemonaceae than with core Resedaceae. This different placements have been cleaved off into their
prevents unneccesary inflation of family names. own families. In APG III (2009), these included Bar-
Cleome L. has been shown to be grossly paraphyletic beuiaceae, Gisekiaceae, Lophiocarpaceae and Steg-
to the other previously recognized genera of Cleo- nospermataceae. Most recent studies (Brockington

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
8 THE ANGIOSPERM PHYLOGENY GROUP

et al., 2009, 2011; Bissinger et al., 2014) have found included in Cornaceae in APG III (2009), have been
that subfamily Rivinoideae of Phytolacaccaeae are shown by molecular studies (Xiang et al., 2011) to
sister to Nyctaginaceae, and we propose here to include Camptothecaceae, Davidiaceae and Mastixi-
accept them at the family level (Petiveriaceae, aceae, which are sister to a clade comprising
including Rivinaceae) to maintain the previous use Hydrostachyaceae, Loasaceae and Hydrangeaceae.

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of family names in this larger clade. The priority of They are therefore widely separated from Cornaceae,
Petiveriaceae C.Agardh 1824 over Rivinaceae and thus Nyssaceae need to be accepted.
C.Agardh 1824 was established by Meisner (1841), In Gentianales, the limits of Gelsemiaceae have
who combined the tribes Rivineae Dumort. and been altered here by the inclusion of Pteleocarpa
Petiverieae Bartl. under Petiveriaceae (Art. 11.5). Oliv., which had previously been considered problem-
The second problematic area in Caryophyllales atic; it had been included by various authors in Borag-
involves Cactaceae and their relationship to the for- inaceae, Cardiopteridaceae and Icacinaceae. It was
mer broadly defined Portulacaceae, the latter shown always an odd element in any family and was thus
to be paraphyletic to Cactaceae. In APG III (2009), sometimes placed in its own family, Pteleocarpaceae
Anacampserotaceae, Montiaceae and Talinaceae (Brummitt, 2011). Refulio-Rodr
ıguez & Olmstead
were accepted, leaving Portulacaceae with only Por- (2014) and Struwe et al. (2014) demonstrated that it
tulaca L. To reduce the number of monogeneric fami- falls as sister to Gelsemiaceae, and we expand that
lies in this clade, Cactaceae could be expanded to family to include it, in agreement with their findings.
include at least Anacampserotaceae and Portula- Ongoing studies in Lamiales have resulted in sev-
caceae, but this was highly unpopular in the online eral unstudied genera being placed, for example
survey (Christenhusz et al., 2015). Sanango Bunting & Duke (previously considered
The third problematic family in Caryophyllales is Loganiaceae) as sister to Gesneriaceae (Perret et al.,
Molluginaceae, which in their broadest sense are poly- 2012), Peltanthera Benth. as sister to Gesneriaceae
phyletic. In APG III (2009), Limeaceae and Lophio- plus Sanango and Calceolariaceae, and Rehmannia
carpaceae were recognized as distinct, and here we Libosch. ex Fisch. & C.A.Mey. as sister to Oroban-
add three additional families (Sch€ aferhoff, M€
uller & chaceae (not in Scrophulariaceae, as previously
Borsch, 2009; Christenhusz et al., 2014): Kewaceae thought; Xia, Wang & Smith, 2009; Refulio-
(with the genus Kewa Christenh., which has been seg- Rodr
ıguez & Olmstead, 2014). The history of investi-
regated from Hypertelis E.Mey. ex Fenzl., the type gating relationships in Lamiales has some similari-
species H. spergulacea E.Mey. ex Fenzl remaining in ties to work on the monocot order Asparagales, in
Molluginaceae), Microteaceae and Macarthuriaceae. which the old family limits were completely altered
These all have distant relationships to each other and by the results of phylogenetic studies. Because no
to the other genera to which they were thought to be previously suggested relationships could be relied
related (Brockington et al., 2009, 2011; Sch€ aferhoff upon in Asparagales, narrow family limits were ini-
et al., 2009; Christin et al., 2011; Christenhusz et al., tially accepted (APG, 1998), but as molecular studies
2014). Further sampling of Molluginaceae is required progressed and more taxa were sampled with more
(Borsch et al., 2015). Finally, Agdestis Moc. & Sess
e molecular data (e.g. Fay et al., 2000; Pires et al.,
ex DC. appears to be sister to Sarcobatus Nees (Sarco- 2006), relationships became clear and larger family
bataceae; Brockington et al., 2011). Agdestidaceae limits could be applied (APG II, 2003; APG III,
may require recognition as a segregate family 2009). These newly circumscribed families were
(Hern
andez-Ledesma et al., 2015), but more data are heterogeneous, but the wider limits as applied in
needed to support this placement or to confirm the APG III (2009) have been generally well accepted
placement in Sarcobataceae. (Wearn et al., 2013). In Lamiales, the old delimita-
tions of Acanthaceae, Lamiaceae, Scrophulariaceae,
etc., were contradicted by molecular studies, and
although we still use many of these names, their cir-
ASTERIDS
cumscriptions are now vastly different. In addition,
In Ericales, Mitrastemonaceae are placed at the end we have seen the proliferation of small families (13),
of the linear sequence for the order because their just as in Asparagales (APG, 1998, had 29 families
exact position in that order is not yet certain. In in Asparagales vs. 12 here). A similar condensation
Barkman et al. (2004), they were sister to Ericaceae. in the number of families recognized in Lamiales
Hardy & Cook (2012) recovered Mitrastemonaceae as may be needed, for the reasons discussed by Chris-
sister to most of the order except the Marcgravi- tenhusz et al. (2015). However, for now, we propose
aceae–Tetrameristaceae–Balsaminaceae clade. the following minor changes: (1) enlarging Gesneri-
Further studies in Cornales have also resulted in a aceae to include Sanango, (2) enlarging Oroban-
change in family circumscriptions. Nyssaceae, chaceae to include Rehmanniaceae and (3)

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
 APG IV 9

acceptance of Mazaceae as separate from Phry- aceae, Lennoaceae, Wellstediaceae, Heliotropiaceae,

maceae, which with Paulowniaceae form a grade Hydrophyllaceae and the Nama L. clade (often
leading to Orobanchaceae (Albach et al., 2009; Xia referred to as ‘Namaceae’, a name that has not been
et al., 2009; Sch€ aferhoff et al., 2010; Fischer, formally published), which have been proposed by
Sch€ aferhoff & M€ uller, 2012). Tentatively, we main- several authors (Weigend & Hilger, 2010; as

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tain Calceolariaceae and Peltanthera as distinct from reviewed by Stevens, 2001). The need to dismember
Gesneriaceae, although more study of these closely a group shown in all analyses to be monophyletic
related taxa is needed. Eventually either Peltanthera was questioned and strongly rejected as an option by
will need to be recognized in its own family or the online survey (Christenhusz et al., 2015).
Peltanthera and Calceolariaceae could be included in Finally, here we treat Vahliaceae, unplaced to
an expanded Gesneriaceae. A formal infrafamilial order in APG III (2009), as another monofamilial
classification of Gesneriaceae would be needed if an order, Vahliales. Vahlia Thunb. was sister to Sola-
expanded circumscription is adopted, but the position nales in Refulio-Rodr
ıguez & Olmstead (2014), but
of many genera in Lamiales is still uncertain (e.g. only in the Bayesian analysis was this position well
Wightia Wall.; Zhou et al., 2014) so further familial supported. In Stull et al. (2015), Vahlia was sister to
realignment is likely in the future. Lamiales but with low support in both Bayesian and
Icacinaceae in their modern, pre-molecular circum- parsimony analyses.
scription comprised c. 54 genera and 400 species, but Recently the Nomenclature Committee for Vascu-
they were known to be non-monophyletic from the lar Plants (NCVP) has approved the conservation of
time of Savolainen et al. (2000). K arehed (2001) Viburnaceae (Applequist, 2013), thus proposing it be
showed the scope of the problem in greater detail, but the correct name for Adoxaceae sensu APG. This out-
the low levels of rbcL gene sequence divergence among come was contrary to the intention of the original
early-diverging lamiids precluded circumscription of proposal (Reveal, 2008), which aimed to maintain
well-supported taxa. Using three plastid genes (ndhF, nomenclatural stability. We therefore do not accept
matK and rbcL), Byng et al. (2014) fared somewhat this decision of the NCVP in the hope that the Gen-
better, but still failed to find a set of well-supported eral Committee will not approve it in its report to
relationships that could serve as the basis of a new the next botanical congress (cf. Applequist, 2013).
classification for these genera/clades. Stull et al. Of the taxa of uncertain position in APG III
(2015) sequenced 50 complete plastid genomes and, (2009), we have now placed Apodanthaceae in Cucur-
combining these with previous data, proposed a reduc- bitales (Filipowicz & Renner, 2010), Cynomoriaceae
tion in the size of Icacinaceae, expansion of Mette- in Saxifragales (see above), Petenaea Lundell in Pete-
niusaceae and recognition of two new-to-APG orders, naeaceae of Huerteales (Christenhusz et al., 2010)
Icacinales (with Icacinaceae and monogeneric and Nicobariodendron Vasudeva Rao & Chakrab. in
Oncothecaceae) and Metteniusiales (with Mettenusi- Celastraceae (Simmons, 2004). We have added sev-
aceae including Emmotaceae and the Apodytes E.Mey. eral genera of uncertain position to the only remain-
ex Arn. clade). Metteniusaceae here comprise 11 gen- ing genus from APG III (2009), Gumillea, hoping
era, expanded from one in APG III (2009), whereas that by drawing attention to these, we increase the
Icacinaceae are reduced to 25 genera (Byng, 2014; likelihood that they will be studied further.
Byng et al., 2014; Stull et al., 2015). Of other families Overall, the changes from APG III (2009) to APG IV
previously segregated from Icacinaceae s.l. by K are- are minimal. Stability is an important aspect of our
hed (2001), Stemonuraceae and Cardiopteridaceae are approach to this classification, and the APG system
retained in Aquifoliales and Pennantiaceae in Apiales, has remained remarkably consistent since its incep-
respectively. This brings resolution and a well-sup- tion. Little remains now that requires attention,
ported conclusion to the investigation of the limits of although reorganizations and changes of familial cir-
orders and families in this part of the lamiids. cumscriptions will continue, particularly in Caryophyl-
Given the ongoing uncertainty over the exact lales, Lamiales and Santalales, for which more data
placement of Boraginaceae s.l., we recognize an are needed to provide a robust picture of generic and
order, Boraginales, to accommodate the family. Refu- familial relationships. The advent of routine whole-
lio-Rodr
ıguez & Olmstead (2014) found Boraginales plastid genome sequencing and nuclear gene sequenc-
as sister to Lamiales, but only in the Bayesian analy- ing should remedy this situation, as it has done for the
sis was this placement well supported. Stull et al. early-diverging lamiids. Of course, new phylogenetic
(2015) placed Boraginales as sister to Gentianales, understanding may necessitate description of new fam-
but again only in their Bayesian analysis was this ilies, as were the cases with Kewaceae, Macarthuri-
well supported. Here we consider Boraginales to aceae, Microteaceae and Petenaeaceae, but this
comprise a single family, Boraginaceae s.l., including appears to be the most likely source of new data that
Boraginaceae s.s., Codonaceae, Cordiaceae, Ehreti- will require future alteration of the APG system.

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
10 THE ANGIOSPERM PHYLOGENY GROUP

ACKNOWLEDGEMENTS Bissinger K, Khoshravesh R, Kotrade JP, Oakley J, Sage

TL, Sage RF, Hartmann HE, Kadereit G. 2014. Gisekia
We gratefully acknowledge the support of Kathy Wil- (Gisekiaceae): phylogenetic relationships, biogeography, and
lis and the Royal Botanic Gardens, Kew, for a work- ecophysiology of a poorly known C4 lineage in the Caryophyl-
shop in September 2015 in which the substance of lales. American Journal of Botany 101: 499–509.

Downloaded from https://academic.oup.com/botlinnean/article/181/1/1/2416499 by National Science & Technology Library user on 03 February 2023
this update was discussed. Borsch T, Hern
andez-Ledesma P, Berendsohn WG,
Flores-Olvera H, Ochoterena H, Zuloaga FO, Mering
S, Kilian N. 2015. An integrative and dynamic approach
for monographing species-rich plant groups – building the
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SUPPORTING INFORMATION
Additional Supporting Information may be found in the online version of this article:
Data S1. Angiosperm phylogeny classification of flowering plants (APG IV) with the families organized alpha-
betically within orders.

Appendix Piperales Bercht. & J.Presl

10 [11]. Saururaceae Rich. ex T.Lestib., nom. cons.

Linear classification of flowering plants (APG IV) 11 [12]. Piperaceae Giseke, nom. cons.
12 [15]. *Aristolochiaceae Juss., nom. cons. (including
*Changed circumscription of a family or families added Asaraceae Vent., Hydnoraceae C.Agardh, nom. cons.,
since APG III (2009). Lactoridaceae Engl., nom. cons.)
†
Orders added since APG III (2009).
Numbers in square brackets are those of LAPG (Haston Magnoliales Juss. ex Bercht. & J.Presl
et al., 2009).
13 [16]. Myristicaceae R.Br., nom. cons.
Amborellales Melikyan et al. 14 [17]. Magnoliaceae Juss., nom. cons.
15 [18]. Degeneriaceae I.W.Bailey & A.C.Sm., nom. cons.
1 [1]. Amborellaceae Pichon, nom. cons. 16 [19]. Himantandraceae Diels, nom. cons.
17 [20]. Eupomatiaceae Orb., nom. cons.
Nymphaeales Salisb. ex Bercht. & J.Presl 18 [21]. Annonaceae Juss., nom. cons.

2 [2]. Hydatellaceae U.Hamann Laurales Juss. ex Bercht. & J.Presl

3 [3]. Cabombaceae Rich. ex A.Rich., nom. cons.
4 [4]. Nymphaeaceae Salisb., nom. cons. 19 [22]. Calycanthaceae Lindl., nom. cons.
20 [23]. Siparunaceae Schodde
Austrobaileyales Takht. ex Reveal 21 [24]. Gomortegaceae Reiche, nom. cons.
22 [25]. Atherospermataceae R.Br.
5 [5]. Austrobaileyaceae Croizat, nom. cons. 23 [26]. Hernandiaceae Blume, nom. cons.
6 [6]. Trimeniaceae Gibbs, nom. cons. 24 [27]. Monimiaceae Juss., nom. cons.
7 [7]. Schisandraceae Blume, nom. cons. 25 [28]. Lauraceae Juss., nom. cons.

MESANGIOSPERMS INDEPENDENT LINEAGE: UNPLACED TO MORE INCLUSIVE

CLADE

MAGNOLIIDS
Chloranthales Mart.
Canellales Cronq.
26 [8]. Chloranthaceae R.Br. ex Sims, nom. cons.
8 [9]. Canellaceae Mart., nom. cons.
9 [10]. Winteraceae R.Br. ex Lindl., nom. cons.

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
 APG IV 15

MONOCOTS
Asparagales Link

Acorales Mart. 61 [62]. Orchidaceae Juss., nom. cons.

62 [63]. Boryaceae M.W.Chase et al .

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27 [29]. Acoraceae Martinov 63 [64]. Blandfordiaceae R.Dahlgren & Clifford
64 [65]. Asteliaceae Dumort.
Alismatales R.Br. ex Bercht. & J.Presl 65 [66]. Lanariaceae H.Huber ex R.Dahlgren
66 [67]. Hypoxidaceae R.Br., nom. cons.
28 [30]. Araceae Juss., nom. cons. 67 [69]. Doryanthaceae R.Dahlgren & Clifford
29 [31]. Tofieldiaceae Takht. 68 [70]. Ixioliriaceae Nakai (as ‘Ixiolirionaceae’; spelling
30 [32]. Alismataceae Vent., nom. cons. corrected)
31 [33]. Butomaceae Mirb., nom. cons. 69 [68]. Tecophilaeaceae Leyb., nom. cons.
32 [34]. Hydrocharitaceae Juss., nom. cons. 70 [71]. Iridaceae Juss., nom. cons.
33 [35]. Scheuchzeriaceae F.Rudolphi, nom. cons. 71 [72]. Xeronemataceae M.W.Chase et al.
34 [36]. Aponogetonaceae Planch., nom. cons. 72 [73]. Asphodelaceae Juss., nom. cons. prop. (including
35 [37]. *Juncaginaceae Rich., nom. cons. Xanthorrhoeaceae Dumort., nom. cons.)
36. *Maundiaceae Nakai 73 [74]. Amaryllidaceae J.St.-Hil., nom. cons.
37 [38]. Zosteraceae Dumort., nom. cons. 74 [75]. Asparagaceae Juss., nom. cons.
38 [39]. Potamogetonaceae Bercht. & J.Presl, nom. cons
39 [40]. Posidoniaceae Vines, nom. cons. Arecales Bromhead
40 [41]. Ruppiaceae Horan., nom. cons.
41 [42]. Cymodoceaceae Vines, nom. cons. 75 [90]. Dasypogonaceae Dumort.
76 [76]. Arecaceae Bercht. & J.Presl, nom. cons. (=
Petrosaviales Takht. Palmae Juss., nom. cons.)

42 [43]. Petrosaviaceae Hutch., nom. cons. Commelinales Mirb. ex Bercht. & J.Presl

Dioscoreales Mart. 77 [77]. Hanguanaceae Airy Shaw

78 [78]. Commelinaceae Mirb., nom. cons.
43 [44]. Nartheciaceae Fr. ex Bjurzon 79 [79]. Philydraceae Link, nom. cons.
44 [45]. ‘Burmanniaceae’ Blume, nom. cons. 80 [80]. Pontederiaceae Kunth, nom. cons.
45 [46]. Dioscoreaceae R.Br., nom. cons. 81 [81]. Haemodoraceae R.Br., nom. cons.

Pandanales R.Br. ex Bercht. & J.Presl Zingiberales Griseb.

46 [47]. Triuridaceae Gardner, nom. cons. 82 [82]. Strelitziaceae Hutch., nom. cons.
47 [48]. Velloziaceae J.Agardh, nom. cons. 83 [83]. Lowiaceae Ridl., nom. cons.
48 [49]. Stemonaceae Caruel, nom. cons. 84 [84]. Heliconiaceae Vines
49 [50]. Cyclanthaceae Poit. ex A.Rich., nom. cons. 85 [85]. Musaceae Juss., nom. cons.
50 [51]. Pandanaceae R.Br., nom. cons. 86 [86]. Cannaceae Juss., nom. cons.
87 [87]. Marantaceae R.Br., nom. cons.
Liliales Perleb 88 [88]. Costaceae Nakai
89 [89]. Zingiberaceae Martinov, nom. cons.
51 [52]. Campynemataceae Dumort.
52 [60]. Corsiaceae Becc., nom. cons. Poales Small
53 [53]. Melanthiaceae Batsch ex Borkh., nom. cons.
54 [54]. Petermanniaceae Hutch, nom. cons. 90 [91]. Typhaceae Juss., nom. cons.
55 [55]. Alstroemeriaceae Dumort., nom. cons. 91 [92]. Bromeliaceae Juss., nom. cons.
56 [56]. Colchicaceae DC., nom. cons. 92 [93]. Rapateaceae Dumort., nom. cons.
57 [57]. Philesiaceae Dumort., nom. cons. 93 [94]. Xyridaceae C.Agardh, nom. cons.
58 [58]. Ripogonaceae Conran & Clifford 94 [95]. Eriocaulaceae Martinov, nom. cons.
59 [59]. Smilacaceae Vent., nom. cons. 95 [96]. Mayacaceae Kunth, nom. cons.
60 [61]. Liliaceae Juss., nom. cons. 96 [97]. Thurniaceae Engl., nom. cons.
97 [98]. Juncaceae Juss., nom. cons.
98 [99]. Cyperaceae Juss., nom. cons.

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
16 THE ANGIOSPERM PHYLOGENY GROUP

99 [102]. *Restionaceae R.Br., nom. cons. (including †

Dilleniales DC. ex Bercht. & J.Presl
Anarthriaceae D.W.Cutler & Airy Shaw,
Centrolepidaceae Endl., nom. cons.) 120 [124]. Dilleniaceae Salisb., nom. cons.
100 [103]. Flagellariaceae Dumort., nom. cons.
101 [104]. Joinvilleaceae Toml. & A.C.Sm.

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102 [105]. Ecdeiocoleaceae D.W.Cutler &
Airy Shaw SUPERROSIDS
103 [106]. Poaceae Barnhart, nom. cons. (= Gramineae
Juss., nom. cons.) Saxifragales Bercht. & J.Presl

121 [125]. Peridiscaceae Kuhlm., nom. cons.

122 [126]. Paeoniaceae Raf., nom. cons.
PROBABLE SISTER OF EUDICOTS
123 [127]. Altingiaceae Lindl., nom. cons.
124 [128]. Hamamelidaceae R.Br., nom. cons.
Ceratophyllales Link 125 [129]. Cercidiphyllaceae Engl., nom. cons.
126 [130]. Daphniphyllaceae M€ ull.Arg., nom. cons.
104 [107]. Ceratophyllaceae Gray, nom. cons. 127 [131]. Iteaceae J.Agardh, nom. cons.
128 [132]. Grossulariaceae DC., nom. cons.
129 [133]. Saxifragaceae Juss., nom. cons.
130 [134]. Crassulaceae J.St.-Hil., nom. cons.
131 [135]. Aphanopetalaceae Doweld
EUDICOTS 132 [136]. Tetracarpaeaceae Nakai
133 [137]. Penthoraceae Rydb. ex Britton, nom. cons.
134 [138]. Haloragaceae R.Br., nom. cons.
Ranunculales Juss. ex Bercht. & J.Presl
135 [139]. Cynomoriaceae Endl. ex Lindl., nom. cons.
105 [108]. Eupteleaceae K.Wilh., nom. cons.
106 [109]. Papaveraceae Juss., nom. cons.
107 [110]. Circaeasteraceae Hutch., nom. cons. ROSIDS
108 [111]. Lardizabalaceae R.Br., nom. cons.
109 [112]. Menispermaceae Juss., nom. cons. Vitales Juss. ex Bercht. & J.Presl
110 [113]. Berberidaceae Juss., nom. cons.
111 [114]. Ranunculaceae Juss., nom. cons. 136 [140]. Vitaceae Juss., nom. cons.

Proteales Juss. ex Bercht. & J.Presl Zygophyllales Link

112 [115]. Sabiaceae Blume, nom. cons. 137 [141]. Krameriaceae Dumort., nom. cons.
113 [116]. Nelumbonaceae A.Rich., nom. cons. 138 [142]. Zygophyllaceae R.Br., nom. cons.
114 [117]. Platanaceae T.Lestib., nom. cons.
115 [118]. Proteaceae Juss., nom. cons. Fabales Bromhead

Trochodendrales Takht. ex Cronq. 139 [143]. Quillajaceae D.Don

140 [144]. Fabaceae Lindl., nom. cons. (= Leguminosae
116 [119]. Trochodendraceae Eichler, nom. cons. Juss., nom. cons.)
141 [145]. Surianaceae Arn., nom. cons.
Buxales Takht. ex Reveal 142 [146]. Polygalaceae Hoffmanns. & Link, nom. cons.

117 [121]. *Buxaceae Dumort., nom. cons. (including Rosales Bercht. & J.Presl
Haptanthaceae C.Nelson)
143 [147]. Rosaceae Juss., nom. cons.
144 [148]. Barbeyaceae Rendle, nom. cons.
145 [149]. Dirachmaceae Hutch.
CORE EUDICOTS
146 [150]. Elaeagnaceae Juss., nom. cons.
147 [151]. Rhamnaceae Juss., nom. cons.
Gunnerales Takht. ex Reveal 148 [152]. Ulmaceae Mirb., nom. cons.
149 [153]. Cannabaceae Martinov, nom. cons.
118 [122]. Myrothamnaceae Nied., nom. cons. 150 [154]. Moraceae Gaudich., nom. cons.
119 [123]. Gunneraceae Meisn., nom. cons. 151 [155]. Urticaceae Juss., nom. cons.

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
 APG IV 17

189 [198]. Putranjivaceae Meisn.

Fagales Engl.
190 [185]. Centroplacaceae Doweld & Reveal
152 [156]. Nothofagaceae Kuprian. 191 [190]. Elatinaceae Dumort., nom. cons.
153 [157]. Fagaceae Dumort., nom. cons. 192 [191]. Malpighiaceae Juss., nom. cons.
193 [192]. Balanopaceae Benth. & Hook.f., nom. cons.

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154 [158]. Myricaceae Rich. ex Kunth, nom. cons.
155 [159]. Juglandaceae DC. ex Perleb, nom. cons. 194 [193]. Trigoniaceae A.Juss., nom. cons.
156 [160]. Casuarinaceae R.Br., nom. cons. 195 [194]. Dichapetalaceae Baill., nom. cons.
157 [161]. Ticodendraceae G
omez-Laur. & L.D.G
omez 196 [195]. Euphroniaceae Marc.-Berti
158 [162]. Betulaceae Gray, nom. cons. 197 [196]. Chrysobalanaceae R.Br., nom. cons.
198 [206]. Humiriaceae A.Juss., nom. cons.
Cucurbitales Juss. ex Bercht. & J.Presl 199 [204]. Achariaceae Harms, nom. cons.
200 [202]. Violaceae Batsch, nom. cons.
159 [163]. *Apodanthaceae Tiegh. ex Takht. 201 [203]. Goupiaceae Miers
160 [164]. Anisophylleaceae Ridl. 202 [199]. Passifloraceae Juss. ex Roussel, nom. cons.
161 [165]. Corynocarpaceae Engl., nom. cons. 203 [200]. Lacistemataceae Mart., nom. cons.
162 [166]. Coriariaceae DC., nom. cons. 204 [201]. Salicaceae Mirb., nom. cons.
163 [167]. Cucurbitaceae Juss., nom. cons. 205 [—]. *Peraceae Klotzsch
164 [168]. Tetramelaceae Airy Shaw 206 [183]. Rafflesiaceae Dumort., nom. cons.
165 [169]. Datiscaceae Dumort., nom. cons. 207 [184]. *Euphorbiaceae Juss., nom. cons.
166 [170]. Begoniaceae C.Agardh, nom. cons. 208 [208]. Linaceae DC. ex Perleb, nom. cons.
209 [209]. *Ixonanthaceae Planch. ex Miq., nom. cons.
[COM-clade; placement uncertain] 210 [188]. Picrodendraceae Small, nom. cons.
211 [189]. Phyllanthaceae Martinov, nom. cons.
Celastrales Link
Geraniales Juss. ex Bercht. & J.Presl
167 [171]. Lepidobotryaceae J.L
eonard, nom. cons.
168 [172]. Celastraceae R.Br., nom. cons. 212 [215]. Geraniaceae Juss., nom. cons.
213 [217]. *Francoaceae A.Juss., nom. cons. (including
Oxalidales Bercht. & J.Presl Bersamaceae Doweld, Greyiaceae Hutch., nom. cons.,
Ledocarpaceae Meyen, Melianthaceae Horan., nom.
169 [173]. Huaceae A.Chev. cons., Rhynchothecaceae A.Juss., Vivianiaceae
170 [174]. Connaraceae R.Br., nom. cons. Klotzsch, nom. cons.)
171 [175]. Oxalidaceae R.Br., nom. cons.
172 [176]. Cunoniaceae R.Br., nom. cons. Myrtales Juss. ex Bercht. & J.Presl
173 [177]. Elaeocarpaceae Juss., nom. cons.
174 [178]. Cephalotaceae Dumort., nom. cons. 214 [218]. Combretaceae R.Br., nom. cons.
175 [179]. Brunelliaceae Engl., nom. cons. 215 [219]. Lythraceae J.St.-Hil., nom. cons.
216 [220]. Onagraceae Juss., nom. cons.
Malpighiales Juss. ex Bercht. & J.Presl 217 [221]. Vochysiaceae A.St.-Hil., nom. cons.
218 [222]. Myrtaceae Juss., nom. cons.
176 [180]. Pandaceae Engl. & Gilg, nom. cons. 219 [223]. Melastomataceae Juss., nom. cons.
177 [207]. *Irvingiaceae Exell & Mendoncßa, nom. cons. 220 [224]. Crypteroniaceae A.DC., nom. cons.
(including Allantospermum Forman) 221 [225]. Alzateaceae S.A.Graham
178 [186]. Ctenolophonaceae Exell & Mendoncßa 222 [226]. Penaeaceae Sweet ex Guill., nom. cons.
179 [181]. Rhizophoraceae Pers., nom. cons.
180 [182]. Erythroxylaceae Kunth, nom. cons. Crossosomatales Takht. ex Reveal
181 [187]. Ochnaceae DC., nom. cons.
182 [212]. Bonnetiaceae L.Beauvis. ex Nakai 223 [227]. Aphloiaceae Takht.
183 [211]. Clusiaceae Lindl., nom. cons. (= Guttiferae 224 [228]. Geissolomataceae A.DC., nom. cons.
Juss., nom. cons.) 225 [229]. Strasburgeriaceae Tiegh., nom. cons.
184 [210]. Calophyllaceae J.Agardh 226 [230]. Staphyleaceae Martinov, nom. cons.
185 [213]. Podostemaceae Rich. ex Kunth, nom. cons. 227 [231]. Guamatelaceae S.H.Oh & D.Potter
186 [214]. Hypericaceae Juss., nom. cons. 228 [232]. Stachyuraceae J.Agardh, nom. cons.
187 [205]. Caryocaraceae Voigt, nom. cons. 229 [233]. Crossosomataceae Engl., nom. cons.
188 [197]. Lophopyxidaceae H.Pfeiff.

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
18 THE ANGIOSPERM PHYLOGENY GROUP

267 [270]. *Resedaceae Martinov, nom. cons. (including

Picramniales Doweld Borthwickiaceae J.X.Su et al ., Stixidaceae Doweld as
‘Stixaceae’, Forchhammeria Liebm.)
Picramniales Doweld 268 [271]. *Capparaceae Juss., nom. cons.
269 [272]. Cleomaceae Bercht. & J.Presl

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230 [234]. Picramniaceae Fernando & Quinn 270 [273]. Brassicaceae Burnett, nom. cons. (= Cruciferae
Juss., nom. cons.)
Huerteales Doweld
SUPERASTERIDS
231 [244]. Gerrardinaceae M.H.Alford
232 [—]. *Petenaeaceae Christenh. et al.
233 [245]. Tapisciaceae Takht. Berberidopsidales Doweld
234 [246]. Dipentodontaceae Merr., nom. cons.
271 [274]. Aextoxicaceae Engl. & Gilg, nom. cons.
Sapindales Juss. ex Bercht. & J.Presl 272 [275]. Berberidopsidaceae Takht.

235 [235]. Biebersteiniaceae Schnizl. Santalales R.Br. ex Bercht. & J.Presl

236 [236]. Nitrariaceae Lindl.
237 [237]. Kirkiaceae Takht. 273 [277]. ‘Olacaceae’ R.Br., nom. cons. [not
238 [238]. Burseraceae Kunth, nom. cons. monophyletic] (including Aptandraceae Miers,
239 [239]. Anacardiaceae R.Br., nom. cons. Coulaceae Tiegh., Erythropalaceae Planch. ex Miq.,
240 [240]. Sapindaceae Juss., nom. cons. (including nom. cons. Octoknemaceae Soler. nom. cons.,
Xanthocerataceae Buerki et al., as ‘Xanthoceraceae’) Strombosiaceae Tiegh., Ximeniaceae Horan.)
241 [241]. Rutaceae Juss., nom. cons. 274 [278]. Opiliaceae Valeton, nom. cons.
242 [242]. Simaroubaceae DC., nom. cons. 275 [276]. Balanophoraceae Rich., nom. cons.
243 [243]. Meliaceae Juss., nom. cons. 276 [279]. ‘Santalaceae’ R.Br., nom. cons. [not
monophyletic if Balanophoraceae are embedded]
Malvales Juss. ex Bercht. & J.Presl (including Amphorogynaceae Nickrent & Der,
Cervantesiaceae Nickrent & Der, Comandraceae
244 [247]. Cytinaceae A.Rich. Nickrent & Der, Nanodeaceae Nickrent & Der,
245 [248]. Muntingiaceae C.Bayer et al. Thesiaceae Vest, Viscaceae Batsch)
246 [249]. Neuradaceae Kostel., nom. cons. 277 [281]. Misodendraceae J.Agardh, nom. cons.
247 [250]. Malvaceae Juss., nom. cons. 278 [282]. Schoepfiaceae Blume
248 [251]. Sphaerosepalaceae Bullock 279 [280]. Loranthaceae Juss., nom. cons.
249 [252]. Thymelaeaceae Juss., nom. cons.
250 [253]. Bixaceae Kunth, nom. cons. Caryophyllales Juss. ex Bercht. & J.Presl
251 [255]. *Cistaceae Juss., nom. cons. (including
Pakaraimaea Maguire & P.S.Ashton) 280 [283]. Frankeniaceae Desv., nom. cons.
252 [254]. Sarcolaenaceae Caruel, nom. cons. 281 [284]. Tamaricaceae Link, nom. cons.
253 [256]. *Dipterocarpaceae Blume, nom. cons. 282 [285]. Plumbaginaceae Juss., nom. cons.
283 [286]. Polygonaceae Juss., nom. cons.
Brassicales Bromhead 284 [287]. Droseraceae Salisb., nom. cons.
285 [288]. Nepenthaceae Dumort, nom. cons.
254 [257]. Akaniaceae Stapf, nom. cons. 286 [289]. Drosophyllaceae Chrtek et al .
255 [258]. Tropaeolaceae Juss. ex DC., nom. cons. 287 [290]. Dioncophyllaceae Airy Shaw, nom. cons.
256 [259]. Moringaceae Martinov, nom. cons. 288 [291]. Ancistrocladaceae Planch. ex Walp., nom. cons.
257 [260]. Caricaceae Dumort., nom. cons. 289 [292]. Rhabdodendraceae Prance
258 [261]. Limnanthaceae R.Br., nom. cons. 290 [293]. Simmondsiaceae Tiegh.
259 [262]. Setchellanthaceae Iltis 291 [294]. Physenaceae Takht.
260 [263]. Koeberliniaceae Engl., nom. cons. 292 [295]. Asteropeiaceae Takht. ex Reveal & Hoogland
261 [264]. Bataceae Mart. ex Perleb, nom. cons. 293 [—]. *Macarthuriaceae Christenh.
262 [265]. Salvadoraceae Lindl., nom. cons. 294 [—]. *Microteaceae Sch€aferhoff & Borsch
263 [266]. Emblingiaceae Airy Shaw 295 [296]. Caryophyllaceae Juss., nom. cons.
264 [267]. Tovariaceae Pax, nom. cons. 296 [297]. Achatocarpaceae Heimerl, nom. cons.
265 [268]. Pentadiplandraceae Hutch. & Dalziel 297 [298]. Amaranthaceae Juss., nom. cons.
266 [269]. Gyrostemonaceae A.Juss., nom. cons. 298 [299]. Stegnospermataceae Nakai
299 [300]. *Limeaceae Shipunov ex Reveal

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 APG IV 19

300 [301]. Lophiocarpaceae Doweld & Reveal †

Icacinales Tiegh.
301 [—]. *Kewaceae Christenh.
302 [302]. Barbeuiaceae Nakai 347 [345]. Oncothecaceae Kobuski ex Airy Shaw
303 [303]. Gisekiaceae Nakai 348 [347]. *Icacinaceae Miers, nom. cons.
304 [304]. Aizoaceae Martinov, nom. cons.

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305 [305]. *Phytolaccaceae R.Br., nom. cons. †
Metteniusales Takht.
306 [—]. *Petiveriaceae C.Agardh (including Rivinaceae
C.Agardh) 349 [346]. *Metteniusaceae H.Karst. ex Schnizl.
307 [306]. Sarcobataceae Behnke
308 [307]. Nyctaginaceae Juss., nom. cons. Garryales Mart.
309 [308]. *Molluginaceae Bartl., nom. cons.
310 [309]. Montiaceae Raf. 350 [348]. Eucommiaceae Engl., nom. cons.
311 [310]. Didiereaceae Radlk., nom. cons. 351 [349]. Garryaceae Lindl., nom. cons.
312 [311]. Basellaceae Raf., nom. cons.
313 [312]. Halophytaceae S.Soriano Gentianales Juss. ex Bercht. & J.Presl
314 [313]. Talinaceae Doweld
315 [314]. Portulacaceae Juss., nom. cons. 352 [350]. Rubiaceae Juss., nom. cons.
316 [315]. Anacampserotaceae Eggli & Nyffeler 353 [351]. Gentianaceae Juss., nom. cons.
317 [316]. Cactaceae Juss., nom. cons. 354 [352]. Loganiaceae R.Br. ex Mart., nom. cons.
355 [353]. *Gelsemiaceae L.Struwe & V.A.Albert
(including Pteleocarpaceae Brummitt)
ASTERIDS 356 [354]. Apocynaceae Juss., nom. cons.

Cornales Link †
Boraginales Juss. ex Bercht. & J.Presl

318 [—].*Nyssaceae Juss. ex Dumort., nom. cons. 357 [356]. Boraginaceae Juss., nom. cons. (including
319 [317]. Hydrostachyaceae Engl., nom. cons. Codonaceae Weigend & Hilger)
320 [321]. Hydrangeaceae Dumort., nom. cons.
†
321 [322]. Loasaceae Juss., nom. cons. Vahliales Doweld
322 [318]. Curtisiaceae Takht.
323 [319]. Grubbiaceae Endl. ex Meisn., nom. cons. 358 [355]. Vahliaceae Dandy
324 [320]. Cornaceae Bercht. & J.Presl, nom. cons.
Solanales Juss. ex Bercht. & J.Presl
Ericales Bercht. & J.Presl
359 [357]. Convolvulaceae Juss., nom. cons.
325 [323]. Balsaminaceae A.Rich., nom. cons. 360 [358]. Solanaceae Juss., nom. cons.
326 [324]. Marcgraviaceae Bercht. & J.Presl, nom. cons. 361 [359]. Montiniaceae Nakai, nom. cons.
327 [325]. Tetrameristaceae Hutch. 362 [360]. Sphenocleaceae T.Baskerv., nom. cons.
328 [326]. Fouquieriaceae DC., nom. cons. 363 [361]. Hydroleaceae R.Br.
329 [327]. Polemoniaceae Juss., nom. cons.
330 [328]. Lecythidaceae A.Rich., nom. cons. Lamiales Bromhead
331 [329]. Sladeniaceae Airy Shaw
332 [330]. Pentaphylacaceae Engl., nom. cons. 364 [362]. Plocospermataceae Hutch.
333 [331]. Sapotaceae Juss., nom. cons. 365 [363]. Carlemanniaceae Airy Shaw
334 [332]. Ebenaceae G€ urke, nom. cons. 366 [364]. Oleaceae Hoffmanns. & Link, nom. cons.
335 [333]. Primulaceae Batsch ex Borkh., nom. cons. 367 [365]. Tetrachondraceae Wettst.
336 [334]. Theaceae Mirb., nom. cons. 368 [366]. Calceolariaceae Olmstead
337 [335]. Symplocaceae Desf., nom. cons. 369 [367]. *Gesneriaceae Rich. & Juss., nom. cons. (note:
338 [336]. Diapensiaceae Lindl., nom. cons. position of Peltanthera Benth. is problematic and here
339 [337]. Styracaceae DC. & Spreng., nom. cons. considered unplaced to family)
340 [338]. Sarraceniaceae Dumort., nom. cons. 370 [368]. Plantaginaceae Juss., nom. cons.
341 [339]. Roridulaceae Martinov, nom. cons. 371 [369]. Scrophulariaceae Juss., nom. cons.
342 [340]. Actinidiaceae Gilg & Werderm., nom. cons. 372 [370]. Stilbaceae Kunth, nom. cons.
343 [341]. Clethraceae Klotzsch, nom. cons. 373 [371]. Linderniaceae Borsch et al .
344 [342]. Cyrillaceae Lindl., nom. cons. 374 [383]. Byblidaceae Domin, nom. cons.
345 [344]. Ericaceae Juss., nom. cons. 375 [384]. Martyniaceae Horan., nom. cons.
346 [343]. Mitrastemonaceae Makino, nom. cons. 376 [372]. Pedaliaceae R.Br., nom. cons.
[placement in order unclear]

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20
20 THE ANGIOSPERM PHYLOGENY GROUP

377 [378]. Acanthaceae Juss., nom. cons.

Paracryphiales Takht. ex Reveal
378 [379]. Bignoniaceae Juss., nom. cons.
379 [377]. Lentibulariaceae Rich., nom. cons. 407 [404]. Paracryphiaceae Airy Shaw
380 [381]. Schlegeliaceae Reveal
381 [380]. Thomandersiaceae Sreem.

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Dipsacales Juss. ex Bercht. & J.Presl
382 [382]. Verbenaceae J.St.Hil., nom. cons.
383 [373]. Lamiaceae Martinov, nom. cons. (= Labiatae 408 [405]. Adoxaceae E.Mey., nom. cons. (= Viburnaceae
Juss., nom. cons.) Raf., nom. cons. prop.)
384 [—].*Mazaceae Reveal 409 [406]. Caprifoliaceae Juss., nom. cons.
385 [374]. *Phrymaceae Schauer, nom. cons.
386 [375]. Paulowniaceae Nakai Apiales Nakai
387 [376]. *Orobanchaceae Vent., nom. cons. (including
Lindenbergiaceae Doweld, Rehmanniaceae Reveal) 410 [407]. Pennantiaceae J.Agardh
411 [408]. Torricelliaceae Hu
Aquifoliales Senft 412 [409]. Griseliniaceae Takht., nom. cons. prop.
413 [410]. Pittosporaceae R.Br., nom. cons.
388 [385]. Stemonuraceae K arehed 414 [411]. Araliaceae Juss., nom. cons.
389 [386]. Cardiopteridaceae Blume, nom. cons. 415 [412]. Myodocarpaceae Doweld
390 [387]. Phyllonomaceae Small 416 [413]. Apiaceae Lindl., nom. cons. (= Umbelliferae
391 [388]. Helwingiaceae Decne. Juss., nom. cons.)
392 [389]. Aquifoliaceae Bercht. & J.Presl, nom. cons.
Incertae sedis
Asterales Link
Atrichodendron Gagnep. (specimen poorly preserved, and
393 [390]. Rousseaceae DC. thus difficult to know to which family it should belong;
394 [391]. Campanulaceae Juss., nom. cons. it is definitely not Solanaceae where it was previously
395 [392]. Pentaphragmataceae J.Agardh, nom. cons. placed, S. Knapp, pers. comm.)
396 [393]. Stylidiaceae R.Br., nom. cons. Coptocheile Hoffmanns. (described in Gesneriaceae and
397 [394]. Alseuosmiaceae Airy Shaw may belong there but may belong elsewhere in
398 [395]. Phellinaceae Takht. Lamiales)
399 [396]. Argophyllaceae Takht. Gumillea Ruiz & Pav. (originally placed in Cunoniaceae,
400 [397]. Menyanthaceae Dumort., nom. cons. where it certainly does not belong; it may be close to
401 [398]. Goodeniaceae R.Br., nom. cons. Picramniales or Huerteales)
402 [399]. Calyceraceae R.Br. ex Rich., nom. cons. Hirania Thulin (described in Sapindales and stated to be
403 [400]. Asteraceae Bercht. & J.Presl, nom. cons. (= related to Diplopeltis, but may belong elsewhere;
Compositae Giseke, nom. cons.) phylogenetic evidence is wanting)
Keithia Spreng. (described in Capparaceae, but may
Escalloniales Link belong elsewhere in Brassicales)
Poilanedora Gagnep. (described in Capparaceae, but does
404 [401]. Escalloniaceae R.Br. ex Dumort., nom. cons. not seem to belong there)
Rumphia L. (only known from illustration)
Bruniales Dumort.

405 [402]. Columelliaceae D.Don, nom. cons.

406 [403]. Bruniaceae R.Br. ex DC., nom. cons.

© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 181, 1–20