Download diversityand classification of flowering plants

180
CHAPTER 6
F
EVOLUTION OF FLOWERING PLANTS
REFERENCES FOR FURTHER STUDY
7
Andrews, H. N. 1961. Studies in Paleobotany.
Wiley, New York.
APG ifi. 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:
105—121.
Crane, P. R., E. M. Friis, and K. Pedersen.
1995. The origin and early diversification of
angiosperms. Nature 374: 27.
Crepet, W. L. 1998. The abominable mystery.
Science 282: 1653—1654.
Cronquist, A. 1981. An integrated system of
classification of flowering plants. Columbia
University Press, New York.
Davies, T. J., T. G. Barraclough, M. W. Chase,
P. 5. Soltis, D. E. Soltis, and V. Savolainen. 2004.
Darwin’s abominable mystery: Insights from
a supertree of the angiosperms. Proceedings
of the National Academy of Sciences of the
United States of America 101: 1904—1909.
Doyle, J. A. 2006. Seed ferns and the origin
of angiosperms. Journal of the Torrey Botani
cal Society 133: 169—209.
Doyle, J. A. 2008. Integrating molecular
phylogenetic and paleobotanical evidence on
origin of the flower. International Journal of
Sciences 169: 816—843.
Plant
Eames, A. J. 1961. Morphology of the angios
perms. McGraw Hill, New York.
Friedman, W. E., and J. H. Williams. 2004.
Developmental evolution of the sexual proces
s in ancient flowering plant lineages. The Plant
16, S119—S 132, Supplement.
Cell
Friedman, W. E., and K. C. Ryerson. 2009.
Reconstructing the ancestral female gametophyte
of angiosperms: insights from Amborella and
other ancient lineages of flowering plants. Ameri
can Journal of Botany 96: 129—143.
Friis, E. M., J. A. Doyle, P. K. Endress, and
Q. Leng. 2003. Archaefructus: Angiosperm precursor or specialized
Plant Science 8: 369—373.
early angiosperm? Trends in
Friis, E. M., K. R. Pedersen, and P. R. Crane.
2000. Reproductive structure and organization
of basal angiosperms from the Early Cretaceous
(Barremian or Aptian) of western Portugal.
International Journal of Plant Sciences 161:
S 169—S 182.
Goethe, J. W. 1790. Versuch die Metamorphos
e der Pflanzen zu erkllien. Ettinger, Gotha,
Germany.
Gould, R. E., and T. Delevoryas. 1977. The
biology of Glossopteris: Evidence from petrifie
d seed-bearing and pollen-bearing organs.
inga 1: 387—399.
Alcher
Jack, T. 2001. Relearning our ABCs: New
twists on an old model. Trends in Plant Scienc
e 6: 310—316.
Jenik, P. D., and V. F. Irish. 2000. Regulation
of cell proliferation patterns by homeotic genes
during Arabidopsis floral development. Deve
opment 127: 1267—1276.
l
Retallack, G., and D. L. Dilcher. 1981. Argum
ents for a glossopterid ancestry of angios
perms. Paleobiology 7: 54—67.
Soltis, D. E., P. S. Soltis, P. K. Endress, and
M. W. Chase. 2005. Angiosperm phylogeny
and evolution. Sinauer, Sunderland, MA.
Stebbins, G. L. 1974. Flowering Plants: Evolut
ion Above the Species Level. Belknap Press
of Harvard University Press, Cambridge,
Sun, G., D. L. Dilcher, S. Zheng, and Z.
MA.
Zhou. 1998. In search of the first flower:
A Jurassic angiosperm, Archaefructus, from
China. Science 282: 1692—1695,
Northeast
Sun, G., Q. Ji, D. L. Dilcher, S. Zheng, K.
C. Nixon, and X. Wang. 2002. Archaefructaceae,
a new basal angiosperm family. Science 296: 899—
Stewart, W. N., and G. W. Rothwell. 1993.
904.
Paleobotany and the evolution of plants. Cambr
idge Univesity Press, New York.
Takhtajan, A. L. 1991. Evolutionary Trends
in Flowering Plants. Columbia University
Press, New York.
Thomas, H. H. 1925. The Caytoniales, a new
group of angiospermous plants from the Jurassi
c rocks of Yorkshire. Philosophical Transactions
the Royal Society of London 213: 299—363.
of
Vejt, B., P.. J. Schmidt, S. Hake, and M.
F Yanofsky. 1993. Maize floral development:
new genes and old mutants. The Plant Cell 5:
Zaiiis, M. J., P. S. Soltis, Y. L. Qiu, E.
1205—1215.
Zimmer, and D. E. Soltis. 2003. Phylogenetic
analyses and perianth evolution in basal angios
Annals of the Missouri Botanical Garden
perms.
90: 129—150.
DIVERSITY AND CLASSIFICATION
OF FLOWERING PLANTS:
AMBORELLALE S, NYMPHAEALE S,
AUSTROBAILEYALES, MAGNOLIIDS,
CERATOPHYLLALES, AND MONOCOTS
INTRODUCTION
]ORANG1OSPE CDES
182
MONOCOLEDONS
200
182
MonocotApomohieS
200
FAMILY DESCRIPTIONS
184
Classification of the Monoco1ed0flS
202
......
185
ACOLES
203
186
Acoraceae
203
187
ALISMATALES
205
187
s.aceae
205
187
Alismataceae
208
PETROSALVIALES
210
187
DIOSCOREALES
210
MAGNOLIIDS
189
Dioscoreaceae
210
LAUPALES
190
Lauraceae
190
187
MAGNOLIALES
192
Annonaceae
Magnoliaceae
192
192
PIPERALES
Aristolochiaceae
Piperaceae
Saururaceae
CERATOPHYLLALES
Ceratophyllaceae
PANDANALES
210
Pandanaceae
210
LILIALES
Liliaceae
213
213
ASPARAGALES
213
Agavaceae
Alliaceae
Amaryllidaceae
Asphodelaceae
Iridaceae
Orchidaceae
Themidaceae
218
220
220
221
224
224
229
192
192
197
197
197
197
(Continued)
© 2010 Elsevier Inc. All right.s reserved.
doi: 10, 1016/B970.012.37438O0.OOOOS
1
181
182
CHAPTER 7
COMMELINIDS
.
230
DASYPOGONACEAE
231
ARECALES
231
Arecaceae (Palmae)
COMMELINALES, ZINGIBERALES, AND POALES
231
232
POALES
Angiospermae
249
Bromeliaceae
Cyperaceae
Eriocaulaceae
Juncaceae
Poaceae (Gramineae)
Restionaceae
Sparganiaceae
Typhaceae
Xyridaceae
250
250
254
254
255
258
262
262
262
COMMELINALES
232
Commelinaceae
Haemodoraceae
Pontederiaceae
235
235
238
ZINGIBERALES
241
REVIEW O.LJESTIONS
265
Musaceae
Strelitziaceae
Zingiberaceae
Cannaceae
241
242
245
245
EXERCISES
266
REFERENCES FOR FURTHER STUDY
266
WEB SITES
274
r MagnoliidS
—
—
c’
c.
—
—
MagnoliophYta
rj
I
Eudicots
T
.—
I
—
—
—
—
—
rj
‘
—
-..
C’,
S
0
—
183
EVOLUTiON AND DIVERSITY OF PLANTS
UNIT 11
DIVERSITY AND CLASSIFICATION OF FLOWERING PLANTS
0
:°
—
.—
cotyledon one
leaf venation parallel
INTRODUCTION
The phylogenetic relationships within the angiosperms has been
and continues to be a field of active research in plant systematics.
Much progress has been made with the use of cladistic method
ology and the incorporation of morphological, anatomical,
embryological, palynological, karyological, chemical, and
molecular data (see Chapters 9—14). The more recent use of mul
tiple gene sequence data has been particularly useful in assessing
higher-level angiosperm relationships. However, the phyloge
netic relationships and classification presented in this chapter
can be viewed as somewhat preliminary, to be further refined
with continued research. For a more precise understanding of
relationships within a particular group, there is no substitute for
consulting the most recent, primary scientific literature.
MAJORANGIOSPERM CL4DES
Portrayal of the relationships of major angiosperm groups is
modeled (with very few exceptions) after the system of the
Angiosperm Phylogeny Group, 2009 (referred to as “APG III
2009”), which supersedes APG 1998 and APG II 2003. The
APG III system is based on published cladistic analyses pri
marily utilizing molecular data (e.g., Chase et al. 1993, 2000;
Graham and Olmstead 2000b; Soltis et al. 1997, 2000, 2007;
Qiu et al. 2000; Zanis et al. 2002) or a combination of mor
phological and molecular data (e.g., Nandi et al. 1998). In the
APG III system, an attempt was made to recognize only those
angiosperm families that are monophyletic. In many cases,
angiosperm families have been redefined from their past, tra
ditional circumscription, either being split into separate groups
(e.g., the traditional “Liliaceae” and “Scrophulariaceae”)
or united into one family (e.g., the Bombacaceae, Malvaceae,
Sterculiaceae, and Tiliaceae united into one family, Malvaceae
s.l.). The APG III system classifies one to several families into
orders (thus, each group having the ending “-ales”), where
strong evidence suggests that the order is monophyletic. It
must be understood, however, that the designated orders are
not comparable evolutionary units and are not indicative of a
hierarchical classification system (see Chapter 2). For exam
ple, a single “order” may be sister to a monophyletic group
containing several orders. The orders can be viewed simply as
convenient placeholders for one or more families that appear
to comprise a monophyletic group with relatively high cer
tainty. Some monophyletic groups containing several orders
are given names, such as Magnoliids, Monocotyledons
(monocots), Commelinids, Eudicotyledons (eudicots), Rosids,
Fabids, Malvids, Asterids, Lamiids, and Campanulids.
The precise interrelationships of the major groups of angio
sperms still show some uncertainty, but recent results have
begun to converge. Figure 7.1 illustrates higher-level phylo
genetic relationships from various analyses that are summa
rized in APG III and modified from Soltis et al. (2007). Note
that some polytomies occur; further research may, in time,
resolve many of these. In particular, the elucidation of the
most basal branches of the flowering plants may yield insight
into early angiosperm evolution and radiation.
As seen in Figure 7.1, the angiosperms can be broadly
delimited into several groups: the Amborellales, Nymphaeales,
Austrobaileyales, Chioranthales, Magnoliids (consisting of
Laurales, Magnoliales, Canellales, and Piperales), monocotyledons (the Monocotyledoneae or monocots), Ceratophyl
lales, and the eudicots. Of these major groups, the current
chapter deals with all but the eudicots, which are covered in
Chapter 8. The Amborellales, Nymphaeales, and Austrobai
leyales are sometimes referred to as “basal” flowering plants
(or the “ANITA” grade) because they include the first clades
that diverged from the common ancestor of the angiosperms.
However, as portrayed in Figure 7.1, it is evident that “basal”
vasculature atactostelic,
vascular cambium absent
sieve tube plastids
pollen tncolpate
or tricolpate-derived
proteiflaceoUS/Cuneate
vessels (secondarily lost in some taxa)
FIGURE 7.1
ifi (2009) and Soltis et al. (2007), with selected apomor
Phylogenetic relationships of major angiosperm clades, after APG
phies (see Chapter 6).
I
angiospermS are a paraphyletic assemblage, not to be recog
nized as a formal taxon. The families within the orders are
listed in Table 7.1 (all except the monocots), Tables 7.2
(“basal” monocots) and 7.3 (commeliflid monocots); eudicot
families are listed in Tables 8.1—8.3 of Chapter 8.
The great bulk of the angiospermS in terms of species diver
sity are contained within the monocots and eudicots. The monocotyledons are a large group, containing approximatelY 22% of
all angiosperms (see later discussion). The eudicots comprise a
very large group, including approximately 75% of all angio
sperms, arid will be treated separately in Chapter 8.
The traditionally defined group “DicotyledOfleae,” the
dicotyledons or dicots, have been defined in the past by their
possession of embryos with two cotyledons. It is now thought
feature
that the possession of two cotyledons is an ancestral
apomorphy for
for the taxa of the flowering plants and not an
delimited
traditionally
any group within. Thus, “dicots” as
and
(all angiospermS other than monocots), are paraphyletic
must be abandoned as a formal taxonomic unit.
In the descriptions in this chapter and in Chapter 8,
exemplars are used for each order or other major group.
context of
The choice of these exemplars is very limited in the
not
the huge diversity of the angiospermS. These treatments are
flower
on
references
fine
designed as a substitute for the many
the end of
ing plant family characteristics (see the references at
of the
this chapter), but are intended as an introduction to some
common or important groups for the beginning student.
Taxa at the traditional rank of family are utilized as pri
described.
mary units; in a few cases subfamilies or tribes are
plant
Only major, general features of commonly encountered
families are presented, with examples cited to show diagnostic
features. More thorough descriptions and illustrations of angio
the
sperm families may be obtained from references cited in
family descriptions and listed at the end of the chapter.
_
182
CHAPTER 7
DIVERSITY AND CLASSIFICATION OF FLOWERING
PLANTS
COMMELINIDS
DASYPOGONACEAE
ARECALES
Arecaceae (Palmae)
COMMELINALES, ZINGIBERALES, AND POALES
COMMELINALES
Commelinaceae
1-Iaemodoraceae
Pontederjaceae
ZINGIBERALES
Musaceae
Streljtzjaceae
Zingiberaceae
Cannaceae
230
231
231
231
232
232
235
235
238
POALES
Bromeljaceae
Cyperaceae
Enocaulaceae
Juncaceae
Poaceae (Gramineae)
Restjonaceae
Sparganiaceae
Typhaceae
Xyridaceae
UNIT [I
Angiospermae
249
250
250
254
254
255
258
262
262
262
183
EVOLUTION AND DIVERSITY OF PLANTS
—
Magnoliophyta
Eudicots
—
o
S
—
r
—
:
•
—
Cl)
rj)
—
-
—
.-
—
•
d
Magnoliids
=
—
c,
Cl,
,
—
c’
Cl)
Cl)
——
—
-..
0
C
0
—
—
.—
..
=C
0
: =
.—
.—
ci) —
—
.‘
:lD-c
241
REVIEW QJESTIONS
241
242
245
245
EXERCISES
INTRODUCTION
The phylogenetic relationships within the angiosperms
has been
and continues to be a field of active research in plant
systematics.
Much progress has been made with the use of cladistic
method
ology and the incorporation of morphological,
anatomical,
embryologica], palynologicai, karyologicai, chemical,
and
molecular data (see Chapters 9—14). The more recent use
of mul
tiple gene sequence data has been particularly
useful in assessing
higher-level angiosperm relationships. However, the
phyloge
netic relationships and classification presented in
this chapter
can be viewed as somewhat preliminary, to be
further refined
with continued research. For a more precise
understanding of
relationships within a particular group, there is no
substitute for
consulting the most recent, primary scientific
literature.
MAJORANGIOSpEI CLADES
Portrayal of the relationships of major angiosperm
groups is
modeled (with very few exceptions) after the
system of the
Angiosperm Phylogeny Group, 2009 (referred
to as “APG III
2009”), which supersedes APG 1998
and APG II 2003. The
APG ifi system is based on published
cladistic analyses pri
marily utilizing molecular data (e.g., Chase
et al. 1993, 2000;
Graham and Olmstead 2000b; Softis et al.
1997, 2000, 2007;
Qiu et al. 2000; Zanis et al. 2002) or a
combination of mor
phological and molecular data (e.g., Nandi
et al. 1998). In the
APG III system, an attempt was made
to recognize only those
angiosperm families that are
monophyletic In many cases,
angiosperm families have been
redefined from their past, tra
ditionad circumscription either being
split into separate groups
(e.g., the traditional “Liliaceae”
and “Scrophulariaceae”)
or united into one family (e.g.,
the Bombacaceae, Malvaceae,
Sterculiaceae, and Tiliaceae united into
one family, Malvaceae
REFERENCES FOR FURTHER STUDY
WEB SITES
265
266
266
274
s.1.). The APG HI system classifies one to several
families into
orders (thus, each group having the ending “-ales”),
where
strong evidence suggests that the order is monophyletic.
It
must be understood, however, that the designated
orders are
not comparable evolutionary units and are not
indicative of a
hierarchical classification system (see Chapter 2). For
exam
ple, a single “order” may be sister to a monophyletic
group
containing several orders. The orders can be viewed
simply as
convenient placeholders for one or more families that
appear
to comprise a monophyletic group with relatively
high cer
tainty. Some monophyletic groups containing several
orders
are given names, such as Magnoliids,
Monocotyledons
(monocots), Commelinids, Eudicotyledons (eudicots),
Rosids,
Fabids, Malvids, Asterids, Lamiids, and Campanulids.
The precise interrelationships of the major groups
of angio
sperms still show some uncertainty, but recent results
have
begun to converge. Figure 7.1 illustrates higher-level
phylo
genetic relationships from various analyses that are
summa
rized in APG III and modified from Soltis et al.
(2007). Note
that some polytomies occur; further research may,
in time,
resolve many of these. In particular, the elucidation
of the
most basal branches of the flowering plants may
yield insight
into early angiosperm evolution and radiation.
As seen in Figure 7.1, the angiosperms can
be broadly
delimited into several groups: the Amborellales,
Nymphaeales,
Austrobaileyales, Chloranthales, Magnoliids (consisting
of
Laurales, Magnoliales, Canellales, and Piperales),
monocotyledons (the Monocotyledoneac or monocots),
Ceratophyl
lales, and the eudicots. Of these major groups,
the current
chapter deals with all but the eudicots, which are
covered in
Chapter 8. The Amborellales, Nymphaeales, and
Austrobaj
leyales are sometimes referred to as “basal” flowering
plants
(or the “ANITA” grade) because they include the
first clades
that diverged from the common ancestor of the
angiosperms.
However, as portrayed in Figure 7.1, it is evident
that “basal”
pollen tricolpate
or tricolpate-derived
vessels (secondarily lost in some taxa)
FIGURE 7.1 Phylogenetic relationships of major angiosperm clades, after APG ifi (2009) and Soltis et al. (2007), with selected apomor
phies (see Chapter 6).
angiosperms are a paraphyletic assemblage, not to be recog
nized as a formal taxon. The families within the orders are
listed in Table 7.1 (all except the monocots), Tables 7.2
(“basal” monocots) and 7.3 (comrnelinid monocots); eudicot
families are listed in Tables 8.1—8.3 of Chapter 8.
The great bulk of the angiosperms in terms of species diver
sity are contained within the monocots and eudicots. The monocotyledons are a large group, containing approximately 22% of
all angiosperms (see later discussion). The eudicots comprise a
very large group, including approximately 75% of all angio
sperms, and will be treated separately in Chapter 8.
The traditionally defmed group “Dicotyledoneae,” the
dicotyledons or dicots, have been defined in the past by their
possession of embryos with two cotyledons. It is now thought
that the possession of two cotyledons is an ancestral feature
for the taxa of the flowering plants and not an apomorphy for
any group within. Thus, “dicots” as traditionally delimited
(all angiosperms other than monocots), are paraphyletic and
must be abandoned as a formal taxonomic unit.
In the descriptions in this chapter and in Chapter 8,
exemplars are used for each order or other major group.
The choice of these exemplars is very limited in the context of
the huge diversity of the angiosperms. These treatments are not
designed as a substitute for the many fine references on flower
ing plant family characteristics (see the references at the end of
this chapter), but are intended as an introduction to some of the
common or important groups for the beginning student.
Taxa at the traditional rank of family are utilized as pri
mary units; in a few cases subfamilies or tribes are described.
Only major, general features of commonly encountered plant
families are presented, with examples cited to show diagnostic
features. More thorough descriptions and illustrations of angio
sperm families may be obtained from references cited in the
family descriptions and listed at the end of the chapter.