229 224 224 220 221 213 220 210 213 213 218 210 210 210 210 205 208 202 203 203 205 200 200
(Continued)
S,
Monoco1ed0flS
the
MONOCOTS
of
PLANTS:
MAGNOLIIDS,
Amaryllidaceae Asphodelaceae Orchidaceae Themidaceae Agavaceae Alliaceae Iridaceae Pandanaceae Liliaceae Alismataceae Dioscoreaceae Acoraceae s.aceae Classification MonocotApomohieS
PANDANALES LILIALES PETROSALVIALES DIOSCOREALES ASPARAGALES MONOCOLEDONS ALISMATALES ACOLES AND NYMPHAEALE
CLASSIFICATION
181
S,
197 197 192 197 192 192 197 189 190 190 192 192 187 186 187 187 187 182 184 185 187 182 7
AND
FLOWERING
CDES
......
reserved.
AMBORELLALE
OF
AUSTROBAILEYALES,
right.s
All
CERATOPHYLLALES,
Inc.
DESCRIPTIONS
Elsevier
1016/B970.012.37438O0.OOOOS
Saururaceae Ceratophyllaceae Piperaceae Annonaceae Magnoliaceae Aristolochiaceae Lauraceae
10,
DIVERSITY
2010
PIPERALES CERATOPHYLLALES LAUPALES MAGNOLIIDS MAGNOLIALES INTRODUCTION ]ORANG1OSPE FAMILY
© doi:
F
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Plant (Barremian
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opment REFERENCES
China. Andrews, APG
Crane, Crepet, Cronquist, the Davies,
Doyle, Annals Doyle,
Eames, Friedman,
Friedman,
Friis, Friis,
Goethe, Gould,
Jack, Jenik,
Retallack, Soltis, Stebbins, Sun,
Sun,
Stewart,
Takhtajan, Thomas, Vejt, Zaiiis,
Sterculiaceae,
or
(e.g.,
ditional
angiosperm APG
angiosperm
phological
Qiu marily
Graham APG
Angiosperm
2009”), modeled
Portrayal
MAJORANGIOSPERM
consulting
relationships
with
can higher-level
netic
molecular
tiple
embryological, Much
ology The
and
INTRODUCTION
182
ZINGIBERALES
ARECALES
COMMELINALES
COMMELINALES,
united
COMMELINIDS
DASYPOGONACEAE
Zingiberaceae
Cannaceae
Musaceae
Strelitziaceae
Pontederiaceae
Haemodoraceae
be
continues
Arecaceae
phylogenetic
III
continued
III
relationships
the
progress
al.
utilizing
and
viewed
circumscription,
which
system,
and
system
into
(with
of
sequence
2000;
data
traditional
and
the
CHAPTER
families
the
families
angiosperm
the
Olmstead Phylogeny
within
(Palmae)
one
and
to
(see
most
molecular
supersedes
has
as
very
research.
molecular
be
relationships
an Zanis incorporation
is
palynological,
relationships
family
Tiliaceae
somewhat
based
and
a
Chapters been
data attempt
.
a
recent,
field
have
few
ZINGIBERALES,
that
particular
“Liliaceae”
et
2000b;
classification
has
Group,
made
either
7
relationships.
(e.g.,
For
of
al.
exceptions)
on
data
been
are
data
APG
been
was
united
active
primary
9—14).
DIVERSITY
2002)
published
preliminary,
a
of
monophyletic.
with
being
(e.g.,
within
the
Soltis
of
more
redefined group,
(e.g.,
2009
made
karyological,
particularly
1998
major
research
into
Bombacaceae, morphological,
The
the
or
and
Nandi
CL4DES
split
scientific
presented
Chase
et precise
the
(referred
to
after
there
AND
a
and
one
use
However,
more
cladistic
angiosperm
al.
combination
recognize
“Scrophulariaceae”) angiosperms
into
from to
in
of
family,
APG
AND
et
1997,
the
is
et
useful
be
POALES
recent
plant
understanding
cladistic
al.
In
separate
no
al.
literature.
chemical,
to
their
in
further
system
analyses
II
1998).
many
substitute
the
2000,
1993,
Malvaceae,
as
CLASSIFICATION
this
systematics.
Malvaceae
in
anatomical,
only
use
2003.
groups
past,
“APG
assessing
phyloge
has
of
method
groups
chapter
of
refined
cases,
of
In
2000;
2007;
those
mor
mul
been
The
tra
pri
the
and
the
for
III
245
245
242
241
241
238
235
235 232
232
231
231
231
230
of
is
However,
that
leyales
Chapter
chapter
(or lales, Laurales,
cotyledons Austrobaileyales,
delimited into resolve
most
that
rized
begun
genetic sperms Fabids,
tainty.
are to
(monocots),
convenient ple,
containing hierarchical
not
must
orders WEB
strong s.l.).
REFERENCES
REVIEW
EXERCISES
POALES
As
The
Xyridaceae
Typhaceae
Sparganiaceae Poaceae
Juncaceae
comprise Restionaceae Eriocaulaceae Bromeliaceae
Cyperaceae
the
diverged
comparable
early
some
a
given
The
basal
in
and
be
SITES OF
seen
single
to
precise
“ANITA”
are
Some
evidence
(thus,
deals
Malvids,
relationships
still
many
APG
8.
understood,
into
converge.
APG
as
O.LJESTIONS
angiosperm (Gramineae)
Magnoliales,
FLOWERING
the
polytomies
sometimes
(the branches
The
several
placeholders
in
names,
portrayed
Commelinids,
classification
show
a
from
“order”
with
monophyletic
several
III
each
interrelationships
of
Figure
III
monophyletic FOR
Monocotyledoneae
Amborellales,
evolutionary
and
grade)
Asterids,
suggests
these.
system
Chioranthales,
the
orders.
all
some
group
Figure
FURTHER
of
such
groups:
modified
from
however,
may
but
referred
evolution
common
7.1,
in
occur;
the
because
In
Of
Canellales,
for
Figure uncertainty,
classifies
the
The
having
system
Lamiids,
flowering
that
Eudicotyledons
be
as
PLANTS
the
various
7.1
particular,
these
groups
one
the
units
group
eudicots,
further
sister
Nymphaeales,
orders
Magnoliids,
from
to
illustrates STUDY
angiosperms
that
the
ancestor
Amborellales,
7.1,
and
of
they
or
as
Magnoliids
the
(see
major
or
the
one
and
analyses
more order
containing
the
and
to
“basal”
Soltis
with
radiation.
it
plants
and
can
monocots),
but
research
include
ending
major
is
the
Chapter
a
to
which
are
designated
Campanulids.
of
evident
monophyletic
groups,
be
families
several
higher-level
relatively
recent
is
Piperales),
elucidation
et
(eudicots),
not
the
may
flowering viewed
Monocotyledons
that
monophyletic.
groups
al.
can
and
the
“-ales”),
are
Nymphaeales,
(consisting
indicative
may,
angiosperms.
several
2).
(2007).
yield
that
results
Ceratophyl
families
are
the
be
first
Austrobai
covered
that
For
orders
simply
high
of
in
summa
“basal” broadly
current
Rosids,
insight
mono-
phylo
plants
appear
angio
of
orders
exam
where
group
time,
Note
have
266
274
265
266
cer
262
262
262
of 258
255
254
254 250
250
into
249
the
are
in of
as
It
a
I
—
FIGURE
phies —
angiospermS
nized
listed
families
(“basal”
sity
cotyledons
all
very
sperms,
possession
that
for
any
The
The
angiosperms
are
the
—
(see
c’
the
group
large
as
in
great
contained
taxa
traditionally
arid
are
monocots) Chapter
possession
a
7.1
Table
—
formal
c.
are
group,
of
within.
listed
will
bulk
of
are
or
Phylogenetic
a
the
(see
6).
7.1
large
be
dicots,
a
—
within
of —
‘
in
including
paraphyletic
flowering
treated
Thus, the and
of
later
(all
Tables
group,
defined
r
two I
with
angiospermS
have the 7.3
C’,
except
The
discussion).
relationships
MagnoliidS
separately
“dicots”
monocots
cotyledons
(commeliflid
8.1—8.3
approximately
two
containing
plants
been
—
families
rj
group
assemblage,
cotyledons.
the
defined
and
as
of
in
in
and
monocots),
The
of
“DicotyledOfleae,”
Chapter
traditionally
terms
Chapter within
is
approximatelY
not
Angiospermae
major
eudicots.
monocots);
an
eudicots
in
75%
an
not
It
ancestral
of
the
angiosperm the
-..
apomorphy
is
0
8.
8.
to
of
past
now
UNIT
Tables
The
orders
comprise
be
all
delimited
eudicot
by
thought
22%
recog
feature
mono-
angio
diver
their
clades,
11
7.2
are
the
for
of
a
cotyledon EVOLUTiON
—
after
rj
must
(all
MagnoliophYta
exemplars
The
the
vasculature
designed
ing vascular
this
common
mary
Only
families
features.
family
sperm
venation
T
In
APG
Taxa
huge
angiospermS
plant
chapter),
choice
proteiflaceoUS/Cuneate one
sieve
vessels
be
the
major,
units;
families
ifi
descriptions
cambium
abandoned
at
diversity
are
family
as
or
More
tube
parallel
(2009)
descriptions
atactostelic,
are
the
of
a
important
AND
in
(secondarily
presented,
general
but
substitute
these
a
thorough
traditional :°
used
characteristics
may
few
and
are
other absent
of
DIVERSITY
0
as
exemplars
the
Soltis
intended
and
cases
features
for
be
groups
a
than
with
for
in
angiospermS.
formal
obtained
descriptions
listed
Eudicots
each
lost
et
rank
the
this
al.
monocots),
examples
—
for
as
of
in
is
many
(see
(2007),
at
order
taxonomic
of
an
the
commonly
chapter
some
very
the
OF
from
family
the
introduction
beginning
pollen
or
.—
fine
and
end
These
limited
with
or
cited
PLANTS
taxa)
references
tricolpate-derived
or
references
are
references
illustrations
and
other
tribes
of
selected
are
tncolpate
unit.
to
encountered
treatments
paraphyletic
the .— -
in
student.
show
in
utilized
to
the
major
are
chapter.
S
at
Chapter
some apomor
cited
on
context
described.
the
diagnostic
—
—
—
of
183
flower
are
group.
as
end
angio
of
in
plant
and
pri
not
I
the
the
of
8, of _
182 CHAPTER 7 DIVERSITY AND CLASSIFICATION OF FLOWERING PLANTS UNIT [I EVOLUTION AND DIVERSITY OF PLANTS 183
COMMELINIDS 230 POALES DASYPOGONACEAE 249 Angiospermae — Magnoliophyta 231 Bromeljaceae 250 ARECALES Cyperaceae 231 250 Eudicots Arecaceae (Palmae) Enocaulaceae 231 254 Magnoliids Cl) Juncaceae rj) 254 r - — COMMELINALES, ZINGIBERALES, AND Poaceae .- — POALES 232 (Gramineae) — Cl) 255 — —— Cl) C COMMELINALES Restjonaceae — — -..Cl, — = 232 258 , 0 Commelinaceae Sparganiaceae 235 262 — — 0 0 C 1-Iaemodoraceae Typhaceae o • c, = 235 262 d — .— Pontederjaceae Xyridaceae — .— .. .— ci) — — .‘ 238 262 S : •— c’ : = ZINGIBERALES :lD-c 241 REVIEW QJESTIONS Musaceae 265 241 EXERCISES Streljtzjaceae 266 242 Zingiberaceae REFERENCES 245 FOR FURTHER STUDY 266 Cannaceae 245 WEB SITES 274
INTRODUCTION s.1.).The APG HI system classifies one to several families into orders (thus, each group having the ending The phylogenetic “-ales”), where relationshipswithin the angiospermshas been strong evidence pollen tricolpate suggests that the order is monophyletic. It and continuesto be a fieldof activeresearch or tricolpate-derived in plant systematics. must be understood, however, that the Much progress designated orders are has been made with the use of cladisticmethod not comparable evolutionary units and are not indicative of a ology and the incorporation of morphological, anatomical, hierarchical classification system (see Chapter 2). For exam embryologica], palynologicai, karyologicai, chemical, and ple, a single “order” may be sister to a monophyletic molecular data (see group Chapters9—14).The more recentuse ofmul containing several orders. The orders can be viewed simply as tiple gene sequencedatahas been particularlyusefulin assessing convenient placeholders for one or higher-level more families that appear angiosperm relationships. However, the to phyloge comprise a monophyletic group with relatively high netic relationships and classification cer presented in this chapter tainty. Some monophyletic groups can be containing several orders viewed as somewhat preliminary, to be further are refined given names, such as Magnoliids, Monocotyledons with continued research. For a more precise vessels (secondarily lost in some taxa) understanding of (monocots), Commelinids, Eudicotyledons (eudicots), relationships within a Rosids, particular group, there is no substitutefor Fabids, Malvids, Asterids, consulting Lamiids, and Campanulids. the most recent, primary scientific literature. The precise interrelationships of the major groups of angio sperms still show some uncertainty, but recent results have FIGURE 7.1 Phylogeneticrelationshipsofmajorangiospermclades,afterAPGifi (2009)andSoltiset al. (2007),withselectedapomor MAJORANGIOSpEI begun to converge. Figure 7.1 illustrates higher-level phylo phies(see Chapter 6). CLADES genetic relationships from various analyses that are summa rized in APG III and modified from Soltis et Portrayal of the relationships of major al. (2007). Note angiosperms are a paraphyletic assemblage, not to be recog (all angiosperms other than monocots), are paraphyletic and angiosperm groups is that some polytomies occur; modeled (with very further research may, in time, nized few exceptions) after the system of the as a formal taxon. The families within the orders are must be abandoned as a formal taxonomic unit. resolve many of these. In particular, the elucidation Angiosperm Phylogeny Group, 2009 (referred of the listed in Table 7.1 (all except the monocots), Tables 7.2 In the descriptions in this chapter and in Chapter 8, to as “APG III most basal branches of the flowering 2009”), which supersedes plants may yield insight (“basal” monocots) and 7.3 (comrnelinid monocots); eudicot exemplars are used for each or group. APG 1998 and APG II 2003. The into early order other major APG ifi angiosperm evolution and radiation. system is based on published cladistic analyses pri families are listed in Tables 8.1—8.3of Chapter 8. The choice of these exemplarsis very limited in the context of As seen in Figure 7.1, the angiosperms can marily utilizing molecular data (e.g., be broadly The great bulk of the angiosperms in terms of species diver the huge diversityof the angiosperms.These treatments are not Chase et al. 1993, 2000; delimited into several Graham and groups: theAmborellales, Nymphaeales, Olmstead 2000b; Softis et al. 1997, 2000, sityare containedwithinthe monocotsand eudicots.The mono- designed as a substitutefor the many fine references on flower 2007; Austrobaileyales, Chloranthales, Magnoliids Qiu et al. 2000; Zanis et al. 2002) (consisting of cotyledons are a large group, containing approximately 22% of ing plant family characteristics(see the references at the end of or a combination of mor Laurales, Magnoliales, phological and Canellales, and Piperales), mono- molecular data (e.g., Nandi et al. 1998). In the all angiosperms (see later discussion).The eudicots comprise a this chapter), but are intendedas an introductionto some of the cotyledons (the Monocotyledoneac or monocots), APG III system, an attempt was made Ceratophyl very large group, including approximately 75% of all angio common or importantgroups for the beginning student. to recognize only those lales, and the eudicots. angiosperm families Of these major groups, the current that are monophyletic In many cases, sperms,and will be treated separatelyin Chapter 8. Taxa at the traditional rank of family are utilized as pri angiosperm chapter deals with all but the eudicots, which families have been redefined are covered in The traditionally defmed group “Dicotyledoneae,” the mary units; in a few cases subfamilies or tribes are described. from their past, tra Chapter 8. The Amborellales, ditionad circumscription either Nymphaeales, and Austrobaj dicotyledons dicots, have been Only plant being split into separate groups leyales or defined in the past by their major, general features of commonly encountered (e.g., are sometimes referred to as “basal” flowering the traditional “Liliaceae” and plants possession of embryos with two cotyledons. It is now thought families are presented, with examples cited to show diagnostic “Scrophulariaceae”) (or the “ANITA” grade) because or united into one family they include the first clades that (e.g., the Bombacaceae, Malvaceae, that the possession of two cotyledons is an ancestral feature features. More thorough descriptions and illustrations of angio Sterculiaceae, diverged from the common ancestor of the angiosperms. and Tiliaceae united into one family, Malvaceae for the taxa of the flowering plants and not an apomorphy for sperm families may be obtained from references cited in the However, as portrayed in Figure 7.1, it is evident that “basal” any group within. Thus, “dicots” as traditionally delimited family descriptions and listed at the end of the chapter.