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Apg Iv
Apg Iv
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 63 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 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
2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016
APG IV 3
Amborellales
Nymphaeales
Austrobaileyales
Angiosperms
Magnoliales
Laurales Magnoliids
Piperales
Canellales
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
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
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
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: xixii) 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 ghems work but in reviews of his articles published
several new orders as a result of studies incorporating in Justs 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 Tieghems 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
APG IV 5
2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016
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
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
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
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
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.
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-Rodrguez & 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- Rodrguez & 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
aceaeTetrameristaceaeBalsaminaceae 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
APG IV 9
2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016
10 THE ANGIOSPERM PHYLOGENY GROUP
2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016
APG IV 11
Chase MW, Zmarzty S, Lled o MD, Wurdack KJ, Swen- Feodorova TA, Voznesenskaya EV, Edwards GE, Roal-
sen SM, Fay MF. 2002. When in doubt, put it in Flacour- son EH. 2010. Biogeographic patterns of diversification
tiaceae: a molecular phylogenetic analysis based on plastid and the origins of C4 in Cleome (Cleomaceae). Systematic
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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.
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.
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APG IV 15
MONOCOTS
Asparagales Link
42 [43]. Petrosaviaceae Hutch., nom. cons. Commelinales Mirb. ex Bercht. & J.Presl
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.
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16 THE ANGIOSPERM PHYLOGENY GROUP
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
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
APG IV 17
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18 THE ANGIOSPERM PHYLOGENY GROUP
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APG IV 19
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]
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