7

Introduction
Most gardeners are familiar with the Latin names for
plants, even if they do not like using them. What
they may also notice on labels in public gardens, as
well as in the RHS Plant Finder, are the family
names, which end in -aceae. The family is the next
main rank above that of genus and is the means to
indicate that genera belonging to the same family
are more similar to each other than to those that
belong in another family. This is part of a botanical
hierarchy used to organise plants into an overall
system of relatedness, generally referred to as a
classification. Man has been classifying plants since
at least Theophrastus (c. 300bc) and our current
system dates back to Linnaeus (1753), who
introduced the concept of the binomial (genus and
species name) as well as the arrangement of plants
according to their flower structure. Both concepts
have dominated our thinking about how we name
and classify plants although, over time, we have been
able to find different ways of collecting information
about plants, and different kinds of data with which
to test our classifications. The most recent of these
techniques has been the analysis of genetic material
(genome) and this has been applied to plants as
much as it has to other kinds of organisms. The
results of analysing many thousands of different
plants have brought about a new understanding of
plant relationships and, consequently, a major
reclassification of plants, and flowering plants
(angiosperms) in particular.
The new classification is known by the acronym
APG (Angiosperm Phylogeny Group) which refers
to the international team of scientists that have been
carrying out the analysis of plant genetic material
over the past 15 to 20 years. There have been three
classifications published (APG, 1998, 2003 and
2009), generally known as APG, APG II and APG
III. While the work has been known about for some
time it is only really over the past five years that its
impact has begun to be felt more widely, with major
institutions such as the Royal Botanic Gardens, Kew
and Edinburgh adopting the APG system for their
herbaria and plant labelling. A recent Plant Network
conference (www.plantnetwork.org/proceeds/
wales2009/summary.htm) showed how the new system
is being taken up in public gardens in the UK
(Latta, 2008; Horticulture Week, 7 August 2009). In
view of this, the RHSs Advisory Committee on
Nomenclature and Taxonomy took the decision that
it was appropriate for the RHS to adopt the
classification as presented in the latest edition of
Mabberleys Plant-book (2008) for use in the RHS
horticultural database, from which the RHS Plant
Finder is produced as well as the labels for plants in
our gardens. This is broadly the same as the
classification presented in the latest version of the
APG classification (APG, 2009) and the differences
are shown in the table with this article.
The importance of classification
A classification is, in the most abstract sense, a
method of organising information. It has at least
two functions: to enable communication, as such
names are a shorthand to indicate relationships, and
to predict, in that in associating entities you should
be able to ascertain the properties of one entity from
its position in the classification. It should also be
possible to incorporate new information. For plants,
as for any other living organisms, that element of
predictiveness means that the basis for the
classification must be genetic relatedness (or shared
descent). Although Linnaeuss most famous
classification, which is predictive, is not based on
relatedness, only shared characters, even Linnaeus
realised its artificiality and drew up a parallel system
based on what he thought was a more natural
arrangement. In the context of classification the
word natural is normally used to mean that it is
based on relatedness. Linnaeus, of course, classified
organisms over 100 years before Darwin published
his book On the Origin of Species by Means of
Natural Selection, which revolutionised the
theoretical basis of classification. It took a while for
the first classifications based on his theory of
evolution to be produced but ever since almost every
classification published represents the classifiers
concept of evolutionary relationships.
The consequence of this is that every scientific
name we use conveys some information about the
plants relationships. The basic unit is the species,
and these are clustered together in genera so that
those that are most closely related belong in the
same genus, those more distantly related will belong
in other genera, which in turn are brought together
in families and so on, in a hierarchy that ultimately
reaches the level of Kingdom (such as Plantae, or
Plants). It is this premise of a plants relationships
being revealed through its name and classification
that is one source for name changes. As relationships
are better understood, so it is necessary to change
the position of the plant and this will generally lead
to a change in name. While this is so, it is also
widely accepted that such classifications are
Plants i n thei r proper places
Plants in their proper places the new
classification of flowering plants
PF_177195.indb 7 4/3/10 10:43:33
8
hypotheses and that the categories in which we
arrange organisms are artificial constructs. In nature
there are no genera or families or orders: some argue
that species are real but as Linnaeus said, nature
does not make leaps, or to put it another way, when
you look at the individuals and populations that
make up plants that you are studying, it is often
hard to see where the divisions lie or whether those
divisions are there at all.
This explanation attempts to show why classifications
matter and, in particular, why the APG classification
and the changes it is making to plant labels and
information about plants matter to gardeners. By
and large the new classification does not affect the
genera although these also are being changed by
the application of molecular methods but does
make significant differences at the levels of family
and above. While molecular data have confirmed
many previously recognised relationships based on
morphological characters, what has been fascinating
is where families not previously thought to be close
together have been found to be closely related. This
has in many cases subsequently brought to light other
features such as plant chemistry and microscopic
characters, generally far less prone to rapid
evolutionary change than those features we have
traditionally relied on, like the flower, that support
the new classification. Molecular data have also
shown that our concepts of what are early evolving
plants and which have a more recent origin need to
change. This too affects the classification as early
evolving plants are customarily placed at the
beginning of a linear sequence of a classification,
while those that appeared later are at the end.
A survey of the new system
A long-standing fundamental division of flowering
plants is based on the number of seed leaves, and
the groups thus distinguished are known as
monocotyledons (one seed-leaf ) and dicotyledons
(two seed-leaves). While the molecular data show
that the monocotyledons are a natural group, they
are found to have arisen from an ancestor with two
seed-leaves and so do not have a separate origin to
the dicots. It also shows that the earliest diverging
flowering plants are dicotyledonous in the
traditional sense of the word. It is of interest that
this relationship is borne out by the structure of the
pollen: the monocots and those dicots that arose
before the monocot-dicot divergence all have pollen
grains with either a single pore or single groove; the
later dicots (or eudicots) all have pollen grains based
on the three-pore or three-groove pattern.
The living flowering plant which diverged first
has been found to be a woody shrub (Amborella)
which is endemic to New Caledonia. While it had
always been regarded as being among the early
arising dicots, it had not been thought to be as
significant in flowering plant evolution as the
molecular data now indicate. It may be something of
a surprise that the water lilies are next in the sequence,
given the apparent complexity and conspicuousness
of their flowers. However, water lilies (Nymphaeaceae)
are beetle-pollinated and it has been observed that
many of the earliest flowering plants are beetle-
pollinated; a reflection of the co-evolution of flowers
and many insect groups. It turns out that beetles are
amongst the earliest insect groups to evolve, and
that flower pollination, and thus flower structure,
has co-evolved with the insect pollinators, as insect
diversity increased rapidly alongside flowering plant
diversity. Also amongst the cluster of early diverging
angiosperms is another group of woody plants and
of these the best known to gardeners are Illicium
(allspice) and Schisandra in the family
Schisandraceae.
Next in the sequence and still before the divergence
of the monocots is a group of predominantly woody,
tropical plants characterised by Magnolia. Those
familiar with previous classifications will be aware
that Magnolia and its allies were thought to be the
earliest flowering plants to arise due to their relatively
unspecialised floral morphology. In addition to
Magnolia, also included in this group are Drimys
(Winteraceae), Aristolochia and Asarum (Aristolochiaceae),
Peperomia (Piperaceae), Chimonanthus (Calycanthaceae),
Laurus and Persea (Lauraceae) and the nutmeg (Myristica,
Myristacaceae).
It is after this point that the monocots and dicots
diverge and the remainder of the dicots are informally
referred to as the eudicots. It is amongst this group
that we start to see the differentiation of perianth
segments into petals and sepals, although this is not
universal and many eudicot flowers are much reduced
or adapted and have lost this feature. The first
eudicot group to arise is that which is represented by
the paradigm of flower morphology, Ranunculus.
Closely related are Berberis and Mahonia (Berberidaceae),
and the poppies (Papaveraceae, which also includes
Corydalis and Dicentra). It is the next group, however,
which probably causes one of the biggest surprises.
Associated together are three families of very different
appearance: another group of water lilies (Nelumbo),
the planes (Platanaceae) and the proteas (Proteaceae).
While there are no obvious features that all three
types of plant share, there are some characteristics
that link the planes and the proteas particularly, most
notably the stipules that surround the stem. Also
among the early diverging eudicots is the box family,
Buxaceae, which in previous classifications was
thought to be closer to the Euphorbiaceae.
This brings us to the group referred to as the
core eudicots amongst which the arrangement of
flower parts in fives is the basic plan, although as
with the differentiation of the perianth, there are
many variations in form where this basic plan is not
Plants i n thei r proper places
PF_177195.indb 8 4/3/10 10:43:34
9
evident. At the bottom of the core eudicots, sister to
all the rest is Gunnera (Gunneraceae), the flower
parts of which are still in twos. Next to the gunneras
comes a large group around the Saxifragaceae
which brings together families which had never
previously been associated, such as Cercidophyllum
(Cercidophyllaceae); Crassula (Crassulaceae); Ribes
(Grossulariaceae); Hamamelis and Corylopsis
(Hamamelidaceae); Itea (Iteaceae) and Paeonia
(Paeoniaceae). This highly diverse group appears to
have evolved rapidly towards the middle of the
Cretaceous and, as yet, no single uniting
characteristic has been identified.
At this point the eudicots divide into two major
lineages, the rosids and the asterids. Broadly this
reflects the organisation of the perianth into those
with separate petals and those where the petals are
fused, at least partially, into a tube. Earlier botanists
referred to these two groups as Polypetalae and
Gamopetalae respectively so it is fascinating to see
this reflected in the results from molecular analysis.
Amongst the rosids there are some more surprises:
the previously unsuspected relationship between the
euphorbias (Euphorbiaceae), the passion flowers
(Passifloraceae), the willows (Salicaceae), the violets
(Violaceae) and the tropical parasite with the largest
solitary flower, Rafflesia. Initially unexpected was the
revelation that the families related to the roses
(Rosaceae) are Cannabis (Cannabaceae), Elaeagnus
(Elaeagnaceae), the elms (Ulmaceae) and the nettles
(Urticaceae). The striking difference in appearance of
the flowers is largely due to wind pollination, which
favours reduced flowers with small petals or no
petals at all, a feature not unknown in the Rosaceae
(e.g. Sanguisorba, Polylepis).
While all these surprises might bring one to
question whether the molecular data are reliable as a
foundation for classification, or that errors may have
crept into the analysis, it should be pointed out that
this work has been carried out on many
representatives of each family and for up to six
different regions of the genome. Although there are
some discrepancies, and these are still the subject of
debate (leading to some plants still being unplaced
in the classification), most of the results are
consistent and therefore give confidence that these
are giving a more accurate picture of relationships.
Another cluster is the group of families which all
share the chemical feature of producing mustard
oils, distinctive for the pungent smell plants in these
families give off, especially if crushed. Although the
presence of mustard oils has been known for some
time, it was not until the molecular data showed
that the plants that produce these oils are related
that the significance of the oils for plant
classification was fully appreciated. This grouping
includes the cabbage family (Brassicaceae), the
capers (Capparaceae), the paw paws (Caricaceae),
Cleome (Cleomaceae) and the nasturtiums
(Tropaeolaceae). Shared chemistry is also to be found
in the group of families related to the pinks
(Caryophyllaceae) which also includes the ice plants
(Aizoaceae), the cacti (Cactaceae), pokeweeds
(Phytolaccaceae) as well as spinach and beets
(Chenopodiaceae); to these have been added, on
molecular evidence, many of the families of
carnivorous plants (e.g. Droseraceae and
Nepenthaceae) as well as the sea heath
(Frankeniaceae) and tamarisk (Tamaricaceae).
In the other main lineage, the asterids, there
are two early diverging groups: one around the
Cornaceae which, unexpectedly, proves to be closer
to the Hydrangeaceae than indicated by floral
morphology; the other, a large cluster around the
Ericaceae which itself now includes the Epacridaceae,
Empetraceae and Pyrolaceae. This cluster contains
families long known to be related but also the phlox
family (Polemoniaceae) and the pitcher plants
(Sarraceniaceae). A contentious issue in APG I and
II has been the boundary between the Primulaceae
and the Myrsinaceae, the latter being largely tropical
shrubs and trees; however, under the current
classification these have been merged into one
family, Primulaceae.
It is amongst the core asterids that some major
families of horticulturally important plants are to
be found and one of the most keenly debated
questions has been the fate of the foxglove family
(Scrophulariaceae). Whereas in many cases the
molecular data have indicated that families might be
merged together, it is quite the reverse in the
Scrophulariaceae where some former members are,
according to APG II and III, placed in new or
different families, such as Calceolariaceae (Calceolaria),
Paulowniaceae (Paulownia) or Phrymaceae (Mimulus).
The greatest upheaval is, perhaps, the transfer of
many genera, such Antirrhinum, Digitalis and Hebe to
the Plantaginaceae, previously a small family of
predominantly wind-pollinated plants. What remains
in the Scrophulariaceae are a few of the original
genera, such as Verbascum and Phygelius, together
with others not originally thought to belong there
including Buddleja and Myoporum. While at first this
may seem an unlikely rearrangement, it should be
remembered that the Scrophulariaceae were never well
defined morphologically.
A similar situation arises with the systematics of
the honeysuckle family (Caprifoliaceae), which in an
earlier version was broken up into a number of
different families. In the latest treatment (APG III),
they have all been combined, together with the
teasel family (Dipsacaceae) and the valerians
(Valerianaceae), into one family, Caprifoliaceae, with
the exception of Sambucus and Viburnum. These
have been referred to the Adoxaceae, a family
originally of small herbaceous woodland plants.
Plants i n thei r proper places
PF_177195.indb 9 4/3/10 10:43:34
10 Plants i n thei r proper places
Some may be surprised by the appearance of the
umbellifers (Apiaceae and the closely related family
Araliaceae) among the asterids alongside Pittosporum
and Griselinia. Also notable is Escallonia,
traditionally considered to be closely related to the
currants (Grossulariaceae) and broadly associated
with the roses, which now comes out as a distinctive
lineage among the asterids.
As mentioned above, the monocots have been
recognised as being a natural group, as defined by
the possession of one seed leaf, in the APG
classification, but changes to relationships within the
group have been proposed. The principal
horticultural groups are the aroids, the palms, the
grasses, bromeliads, gingers, orchids and the diverse
cluster of bulbous, cormous or rhizomatous
monocot plants. These latter have been treated in
many different ways, from a broadly circumscribed
Liliaceae to a large number of much smaller families
(Hyacinthaceae, Convallariaceae, etc.). Molecular
data have helped to resolve many of these
uncertainties and, as with the dicots, have pointed
out a number of more unexpected groupings.
The earliest diverging monocot lineage has been
found to be that leading to Acorus (sweet flag),
previously thought to be a member of the Araceae
(Arum family) and it is perhaps interesting to
observe that among the early diverging monocots,
the majority are aquatic. Other distinctive groups of
monocots such as the palms (Arecaceae) and the
gingers (Zingiberaceae) are still recognised, although
the pineapple family (Bromeliaceae) has been
revealed to be much more closely related to the
grasses (Poaceae) than previously thought.
Molecular data have helped greatly to improve
our understanding of the bulbous petaloid
monocotyledons and given greater significance to a
character, the nature of the seed coat, which had not
formerly been seen to be of importance. In this
group of plants there are two types of seed coat; one
type has a dark, non-cellular covering, the other has
a pale, cellular covering. This distinction neatly fits
with the molecular data, and consequently two main
groups are recognised: the asparagoids (Asparagales)
with dark-coated seeds, and the lilioids (Liliales)
with pale-coated seeds. The Liliaceae is now
circumscribed as a much smaller group of genera,
including Lilium, Tulipa, Fritillaria, Erythronium
and Tricyrtis; while Disporum, Uvularia and
Colchicum are now in the Colchicaceae.
The other main cluster, the asparagoids, includes
an unchanged Iridaceae and Amaryllidaceae, which
may be expanded to include Agapanthus and the
Alliaceae (onions). Many of the genera formerly
included in Liliaceae are now placed in an enlarged
Asparagaceae, such as Convallaria, Ornithogalum,
Scilla, Agave, Cordyline, Polygonatum and Ruscus.
Phormium and Hemerocallis come together in the
Hemerocallidaceae. The orchids, however, long held
to be an isolated group among the monocots, are
now shown to belong among the asparagoids,
reflecting the over-emphasis given in previous
classifications to conspicuous floral features of this
highly adapted group of plants.
Further research
As has been observed on a number of points in this
piece, the more startling rearrangements revealed by
molecular analysis often lack the shared characteristics
that would enable recognition by more traditional
means. The task now is to re-examine the many kinds
of data available to characterise plants to see which
one or ones fit the molecular data. These will vary
from group to group as evolution is driven by
different pressures over time. Further, there are still
uncertainties in the position of some groups, such as
the Boraginaceae, and these questions are still being
worked on, so further changes can be expected,
although these will be relatively minor. As this
research continues and new information comes to
light, these are incorporated in the Angiosperm
Phylogeny Website (www.mobot.org/MOBOT/
Research/APweb/welcome.html). This is an invaluable
resource for anyone interested in finding out more.
For those that prefer their information in print, the
standard reference is the third edition of Mabberleys
Plant-book but the updated edition of Heywood et al.
(Flowering Plants of the World) is also based on APG,
although it diverges significantly in the treatment of
certain families.
Dr John David
RHS Chief Scientist
References
Angiosperm Phylogeny Group (1998). An Ordinal
Classification for the Families of Flowering Plants.
Annals of the Missouri Botanical Garden 85:
531553.
Angiosperm Phylogeny Group (2003). An Update of
the Angiosperm Phylogeny Group Classification for
the Orders and Families of Flowering Plants: APG
II. Botanical Journal of the Linnean Society 141:
399436.
Angiosperm Phylogeny Group (2009). An Update of
the Angiosperm Phylogeny Group Classification for
the Orders and Families of Flowering Plants: APG
III. Botanical Journal of the Linnean Society 161:
105121.
Heywood, V.H., Brummitt, R.K., Culham, A. &
Seberg, O. (2007). Flowering Plant Families of the
World. Kew, RBG Kew.
Latta, J. (2008). Changing to APG II Theory Put
into Practice. Sibbaldia 6: 133153.
Mabberley, D.J. (2008). Mabberleys Plant-book.
Third Edition. Cambridge, CUP.
PF_177195.indb 10 4/3/10 10:43:34
11 Plants i n thei r proper places
Only those families with one or more representative
genera in cultivation are given. Groups in square
brackets are ones for which there are no
representatives in UK horticulture.
Subclass MAGNOLIIDAE
[Superorder Amborellanae]
Superorder Nymphaeanae
Order Nymphaeales
Family Nymphaeaceae (Nymphaea)
Superorder Austrobaileyanae
Order Austrobaileyales
Family Schizandraceae (Schisandra, Illicium)
Order Chloranthales
Family Chloranthaceae (Chloranthus)
Superorder Magnolianae
Order Canellales
Family Winteraceae (Drimys, Pseudowintera)
Order Piperales
Family Aristolochiaceae (Aristolochia, Asarum)
Family Saururaceae (Saururus)
Order Magnoliales
Family Annonaceae (Asimina)
Family Magnoliaceae (Magnolia, Liriodendron)
Order Laurales
Family Atherospermataceae (Atherosperma,
Laurelia)
Family Calycanthaceae (Chimonanthus,
Calycanthus)
Family Lauraceae (Laurus, Lindera, Neolitsea,
Sassafras)
Monocots
Superorder Lilianae
Order Alismatales
Family Acoraceae
1
(Acorus)
Family Araceae (Arum, Dracunculus, Arisaema,
Zantedeschia, Lysichiton)
[Order Petrosaviales]
Order Dioscoreales
Family Dioscoreaceae (Dioscorea, Tacca)
Order Pandanales
Family Pandanaceae (Pandanus)
Order Liliales
Family Alstroemeriacae (Alstroemeria, Bomarea)
Family Colchicaceae (Colchicum, Disporum,
Uvularia, Gloriosa)
1 Treated as a separate order, Acorales in APG III
Family Liliaceae (Lilium, Erythronium,
Fritillaria, Tulipa)
Family Melanthiaceae (Veratrum, Paris, Trillium)
Family Philesiaceae (Philesia, Lapageria)
Family Smilacaceae (Smilax)
Order Asparagales
Family Amaryllidaceae (Amaryllis, Narcissus,
Galanthus, Sternbergia, Crinum)
Family Alliaceae
2
(Allium, Tulbaghia,
Nectaroscordum)
Family Agapanthaceae
2
(Agapanthus)
Family Asparagaceae (Agave, Hosta, Yucca,
Cordyline, Convallaria, Polygonatum, Ruscus,
Asparagus, Triteleia, Scilla, Hyacinthus,
Ornithogalum, Eucomis, Dianella)
Family Asteliaceae (Astelia)
Family Hypoxidaceae (Rhodohypoxis)
Family Iridaceae (Iris, Gladiolus, Moraea,
Sisyrinchium, Crocus, Watsonia, Crocosmia,
Dierama)
Family Ixoliriaceae (Ixolirion)
Family Orchidaceae (Bletilla, Cypripedium,
Dactylorhiza, Pleione, Dendrobium)
Family Tecophilaeaceae (Tecophilaea)
Family Hemerocallidaceae
3
(Hemerocallis,
Phormium)
Family Asphodelaceae
3
(Asphodeline,
Kniphofia, Eremurus, Aloe, Gasteria)
Family Xanthorrhoeaceae (Xanthorrhoea)
[Order Dasypogonales]
Order Arecales
Family Arecaceae (Palmae) (Trachycarpus,
Phoenix)
Order Poales
Family Bromeliaceae (Dyckia, Fascicularia,
Puya)
Family Cyperaceae (Cyperus)
Family Restionaceae (Thamnochortus, Elegia)
Family Poaceae (Poa, Miscanthus, Arundo,
Phyllostachys, Cortaderia, Stipa, Zea)
Order Commelinales
Family Commelinaceae (Tradescantia,
Commelina)
Family Pontederiaceae (Pontederia)
Family Haemadoraceae (Anigozanthus)
Order Zingiberales
Family Musaceae (Musa)
Family Strelitziaceae (Strelitzia)
2 Both families are included in Amaryllidaceae in APG III
3 Both families are included in Xanthorrhoeaceae in APG
III
Classification of a selection of plants found in UK
horticulture according to the arrangement in
Mabberleys Plant-book, Ed. 3 (2008)
PF_177195.indb 11 4/3/10 10:43:34
12 Plants i n thei r proper places
Family Zingiberaceae (Cautleya, Hedychium,
Roscoea)
Family Cannaceae (Canna)
Eudicots
Superorder Ceratophyllanae
Order Ceratophyllales
Family Ceratophyllaceae (Ceratophyllum)
Superorder Ranunculanae
Order Ranunculales
Family Eupteleaceae (Euptelea)
Family Lardizabalaceae (Akebia, Decaisnea,
Holboellia)
Family Menispermaceae (Menispermum)
Family Berberidaceae (Berberis, Nandina,
Podophyllum, Epimedium)
Family Ranunculaceae (Clematis, Anemone,
Adonis, Delphinium, Aquilegia, Helleborus,
Glaucidium)
Family Papaveraceae (Papaver, Corydalis,
Romneya, Macleaya, Meconopsis, Dicentra)
Unplaced Order Sabiales
4
Family Sabiaceae (Meliosma)
Superorder Proteanae
Order Proteales
Family Proteaceae (Protea, Banksia, Grevillea,
Hakea, Telopea, Embothrium)
Family Platanaceae (Platanus)
Unplaced Order Trochodendrales
4
Family Trochodendraceae (Trochodendron,
Tetracentron)
Superorder Buxanae
Order Buxales
Family Buxaceae (Buxus, Sarcococca, Pachysandra)
Core Eudicots
Superorder Myrothamnanae
Order Gunnerales
Family Gunneraceae (Gunnera)
Superorder Berberidopsidanae
Order Berberidopsidales
Family Berberidopsidaceae (Berberidopsis,
Aextoxicon)
Unplaced order Dilleniales
5
Family Dilleniaceae (Hibbertia)
4 The position of these two orders with respect to the
Proteanae and Buxanae is still unresolved and hence they
are treated as unplaced
5 Order Dilleniales not recognised in APG III; the family is
left as unplaced
Superorder Caryophyllanae
Order Caryophyllales
Family Nepenthaceae (Nepenthes)
Family Droseraceae (Dionaea, Drosera)
Family Tamaricaceae (Tamarix)
Family Frankeniaceae (Frankenia)
Family Plumbaginaceae (Plumbago,
Ceratostigma, Armeria, Limonium)
Family Polygonaceae (Polygonum, Persicaria,
Rheum)
Family Caryophyllaceae (Dianthus, Lychnis,
Gypsophila)
Family Amaranthaceae (Amaranthus, Beta,
Chenopodium)
Family Portulacaceae (Portulaca, Lewisia,
Calandrinia)
Family Cactaceae (Opuntia, Cereus)
Family Aizoaceae (Carpobrotus, Delosperma,
Ruschia)
Family Phytolaccaceae (Phytolacca, Ercilla)
Family Nyctaginaceae (Mirabilis)
Superorder Santalanae
Order Santalales
Family Loranthaceae (Viscum)
Order Saxifragales
6
Family Cercidophyllaceae (Cercidophyllum)
Family Crassulaceae (Crassula, Kalanchoe,
Sedum, Sempervivum, Echeveria)
Family Daphniphyllaceae (Daphniphyllum)
Family Grossulariaceae (Ribes)
Family Haloragaceae (Haloragis)
Family Hamamelidaceae (Hamamelis, Parrotia,
Corylopsis, Liquidambar
7
)
Family Iteaceae (Itea)
Family Paeoniaceae (Paeonia)
Family Saxifragaceae (Saxifraga, Astilbe, Bergenia,
Darmera, Heuchera, Rodgersia, Tiarella)
Superorder Rosanae
Order Vitales
Family Vitaceae (Vitis, Parthenocissus)
Order Crossosomatales
Family Staphyleaceae (Staphylaea)
Family Stachyuraceae (Stachyurus)
Order Geraniales
8
Family Melianthaceae (Melianthus,
Greyia)
Family Francoaceae
9
(Francoa)
Family Geraniaceae (Geranium, Pelargonium,
Erodium)
Order Myrtales
8
6 Treated as unplaced in APG III, i.e. not assigned to a
Superorder
7 Recognised in APG III in the separate family Altingiaceae
8 APG III has these orders in the Malvids Eurosids II
9 Included in Melianthaceae in APG III
PF_177195.indb 12 4/3/10 10:43:34
13 Plants i n thei r proper places
Family Lythraceae (Lythrum, Punica,
Lagerstroemia, Cuphea, Heimia)
Family Onagraceae (Fuchsia, Clarkia,
Oenothera, Gaura)
Family Myrtaceae (Myrtus, Eucalyptus,
Callistemon, Luma, Acca, Leptospermum)
Family Melastomataceae (Tibouchina,
Osbeckia, Heterocentron)
Fabids Eurosids I
Order Zygophyllales
Family Zygophyllaceae (Peganum)
Order Celastrales
Family Celastraceae (Euonymus, Maytenus,
Parnassia)
Order Malpighiales
Family Salicaceae
10
(Salix, Azara, Idesia,
Poliothyrsis, Populus)
Family Violaceae (Viola, Melicytus)
Family Passifloraceae (Passiflora)
Family Euphorbiaceae (Euphorbia, Ricinus,
Codiaeum, Acalypha, Croton, Mallotus)
Family Hypericaceae (Hypericum)
Family Linaceae (Linum)
Order Oxalidales
Family Oxalidaceae (Oxalis, Biophytum)
Family Cunoniaceae (Eucryphia, Weinmannia)
Family Elaeocarpaceae (Crinodendron, Aristotelia)
Order Fabales
Family Quillajaceae (Quillaja)
Family Fabaceae (Acacia, Cercis, Gleditsia,
Caesalpinia, Sophora, Baptisia, Cytisus,
Genista, Lupinus, Indigofera, Wisteria,
Phaseolus, Robinia, Coronilla, Clianthus,
Trifolium, Lathyrus, Pisum)
Family Polygalaceae (Polygala)
Order Rosales
Family Rosaceae (Rosa, Potentilla, Geum,
Alchemilla, Acaena, Filipendula, Exochorda,
Spiraea, Kerria, Dryas, Rubus, Prunus,
Chaenomeles, Malus, Sorbus, Cotoneaster,
Pyracantha)
Family Elaeagnaceae (Elaeagnus, Hippophae)
Family Rhamnaceae (Rhamnus, Ceanothus,
Colletia)
Family Ulmaceae (Ulmus, Celtis, Zelkova)
Family Cannabaceae (Cannabis, Humulus)
Family Moraceae (Morus, Ficus, Broussonetia)
Family Urticaceae (Urtica, Boehmeria)
Order Cucurbitales
Family Corynocarpaceae (Corynocarpus)
Family Coriariaceae (Coriaria)
Family Cucurbitaceae (Cucumis, Cucurbita,
Bryonia, Ecballium, Citrullus)
Family Datiscaceae (Datisca)
10 Includes a number of genera formerly assigned to the
Flacourtiaceae
Family Begoniaceae (Begonia)
Order Fagales
Family Nothofagaceae
11
(Nothofagus)
Family Fagaceae (Fagus, Quercus, Castanea,
Lithocarpus)
Family Myricaceae (Myrica, Morella, Comptonia)
Family Betulaceae (Corylus, Betula, Carpinus,
Alnus)
Family Casuarinaceae (Allocasuarina)
Family Juglandaceae (Juglans, Carya,
Pterocarya)
Malvids Eurosids II
[Order Huerteales]
[Order Picramniales]
Order Brassicales
Family Tropaeolaceae (Tropaeolum)
Family Limnanthaceae (Limnanthes)
Family Resedaceae (Reseda)
Family Capparaceae (Capparis)
Family Cleomaceae (Cleome)
Family Brassicaceae (Brassica, Erysimum,
Aubrieta, Matthiola, Lunaria, Iberis,
Raphanus, Crambe)
Order Malvales
Family Malvaceae (Tilia, Lavatera,
Fremontodendron, Hoheria, Althaea, Alcea,
Hibiscus, Abutilon)
Family Thymelaeaceae (Daphne, Edgeworthia,
Wikstroemia)
Family Cistaceae (Cistus, Halimum,
Helianthemum)
Order Sapindales
Family Sapindaceae (Acer, Aesculus, Dodonaea,
Koelreuteria)
Family Simourabaceae (Ailanthus, Picrasma)
Family Anacardiaceae (Rhus, Cotinus, Schinus)
Family Rutaceae (Citrus, Choisya, Skimmia,
Phellodendron, Dictamnus, Ruta, Cneorum)
Superorder Asteranae
Order Cornales
Family Cornaceae (Cornus, Alangium)
Family Nyssaceae (Nyssa, Davidia)
Family Hydrangeaceae (Hydrangea, Deutzia,
Philadelphus, Carpenteria, Kirengashoma)
Family Loasaceae (Loasa, Mentzelia)
Order Ericales
Family Balsaminaceae (Impatiens)
Family Polemoniaceae (Phlox, Polemonium,
Cantua, Cobaea, Gilia)
Family Theaceae (Camellia, Stewartia)
Family Pentaphylacaceae (Cleyera, Eurya)
Family Ebenaceae (Diospyros)
Family Primulaceae (Primula, Cyclamen,
Dodecatheon, Lysimachia, Myrsine)
11 Previously included in the Fagaceae
PF_177195.indb 13 4/3/10 10:43:34
14 Plants i n thei r proper places
Family Styracaceae (Styrax, Halesia,
Pterostyrax)
Family Diapensiaceae (Diapensia, Galax,
Shortia)
Family Actinidiaceae (Actinidia,
Clematoclethra, Saurauia)
Family Sarraceniaceae (Sarracenia)
Family Clethraceae (Clethra)
Family Ericaceae
12
(Erica, Calluna, Cassiope,
Kalmia, Rhododendron, Vaccinium, Pyrola,
Empetrum, Cyathodes, Richea)
Lamiids Euasterids I
Family Boraginaceae
13
(Borago, Echium,
Nemophila, Anchusa, Lithodora, Omphalodes,
Pulmonaria, Phacelia)
Order Garryales
Family Eucommiaceae (Eucommia)
Family Garryaceae (Aucuba, Garrya)
Order Gentianales
Family Rubiaceae (Coprosma, Bouvardia,
Luculia, Asperula, Phuopsis)
Family Gentianaceae (Gentiana, Eustoma,
Exacum, Centaurium)
Family Apocynaceae
14
(Nerium,
Trachelospermum, Vinca, Asclepias, Hoya,
Dregea, Stapelia)
Order Solanales
Family Solanaceae (Solanum, Lycium,
Cestrum, Nicotiana, Datura, Petunia,
Salpiglossis)
Family Convolvulaceae (Convolvulus,
Ipomoea)
Order Lamiales
Family Oleaceae (Fraxinus, Jasminum, Olea,
Osmanthus, Syringa, Ligustrum)
Family Gesneriaceae (Haberlea, Ramonda,
Mitraria, Achimenes, Streptocarpus)
Family Calceolariaceae (Calceolaria, Jovellana)
Family Scrophulariaceae (Verbascum,
Phygelius, Diascia, Nemesia, Buddleja,
Myoporum)
Family Acanthaceae (Acanthus, Justicia,
Strobilanthes)
Family Verbenaceae (Aloysia, Verbena,
Rhaphithamnus)
Family Bignoniaceae (Catalpa, Eccremocarpus,
Campsis, Incarvillea)
Family Lamiaceae
15
(Lamium, Clerodendrum,
Vitex, Callicarpa, Caryopteris, Phlomis,
Salvia, Mentha, Thymus, Plectranthus,
Stachys)
12 Including Empetraceae and Epacridaceae
13 Family is unplaced (unassigned to an order) in both
Mabberley & APG III; also includes Hydrophyllaceae
14 Including Asclepiadaceae
15 Including some genera formerly in the Verbenaceae
Family Phrymaceae (Mimulus)
Family Paulowniaceae (Paulownia)
Family Orobanchaceae (Castilleja, Lathraea)
Family Plantaginaceae
16
(Digitalis,
Antirrhinum, Penstemon, Hebe, Rehmannia,
Bacopa)
Campanulids Euasterids II
Desfontaniaceae
17
(Desfontania)
Escalloniaceae
18
(Escallonia)
Order Aquifoliales
Family Helwingiaceae (Helwingia)
Family Aquifoliaceae (Ilex)
Order Apiales
Family Griseliniaceae (Griselinia)
Family Araliaceae (Aralia, Schefflera,
Pseudopanax, Hedera, Hydrocotyle)
Family Apiaceae (Astrantia, Eryngium, Daucus,
Bupleurum, Angelica, Foeniculum)
Family Pittosporaceae (Billardieria,
Pittosporum)
Order Dipsacales
Family Adoxaceae (Adoxa, Viburnum,
Sambucus)
Family Caprifoliaceae
19
(Lonicera, Abelia,
Diervilla, Linnaea, Knautia, Scabiosa,
Valeriana, Centranthus, Morina)
Order Asterales
Family Rousseaceae (Carpodetus)
Family Campanulaceae
20
(Campanula,
Platycodon, Adenophora, Codonopsis,
Lobelia)
Family Argophyllaceae (Corokia)
Family Menyanthaceae (Menyanthes)
Family Goodeniaceae (Scaevola)
Family Asteraceae (Aster, Echinops,
Eupatorium, Centaurea, Senecio, Dahlia,
Helichrysum, Osteospermum, Bellis,
Erigeron, Chrysanthemum, Helianthus,
Mutisia, Olearia)
16 Including Globulariaceae
17 Treated as unplaced family by Mabberley, but included
in the Columelliaceae, in the order Bruniales (not
recognised by Mabberley) in APG III
18 Treated as unplaced family by Mabberley, but placed in
the order Escalloniales (not recognised by Mabberley) in
APG III
19 Including Dipsaceae, Valerianaceae and Morinaceae
20 Including Lobeliaceae
PF_177195.indb 14 4/3/10 10:43:34