Plant systematics : an integrated approach

714 Pages·2010·18.49 MB·English

by  Gurcharan Singh

Plant Systematics
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Third edition
Gurcharan Singh
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Chapter 6
Preface
This third edition of integrated information on Plant Systematics has largely been influenced
by the developments of the first few years of twenty first century. Past two decades have
seen development of new tools of biotechnology, vigorous utilization of molecular data in
understanding phylogeny, and redefining affinities and arrangements of plant groups. Recent
years have also seen disappearance of gaps between numerical and cladistic methodologies,
and integration of former into the latter for complete understanding of phylogenetic
relationships. These trends have largely influenced the combination of numerical and
cladistic methods under one chapter, and enlarged discussion on Molecular Systematics,
discussing new concepts, tools and recent achievements. New chapters on Pteridophytes
and Gymnosperms have been added for complete understanding of systematics of vascular
plants.
It is being increasingly realized that actual photographs of plants and plant parts enable
better understanding of taxonomic information, the trend usefully exploited by recent
publications by Simpson (2006) and Judd et al. (3rd ed., 2008). The present edition
incorporates more than 500 colour photographs of plants from diverse families of plants.
High-resolution images of these as also the additional plants have been provided in the
CD-ROM being supplied along with the book, latter including 772 photographs. This has
largely been possible through the kind courtesy of my son Manpreet Singh and daughter-
in-law Komal, who sponsored my recent visit to California, and provided me the opportunity
to visit and photograph temperate plants in and around California. The book as such contains
images of both tropical plants (largely from Delhi), temperate American plants and plants
from other parts of the World growing in the Botanical Gardens of University of California
and San Francisco Botanical Garden. I wish to record the help rendered by the members of
TAXACOM in the identification of some of the American plants.
The focus of the present edition has been to further consolidate the information on the
principles of plant systematics, include detailed discussion on all major systems of
classification, and significantly, also include discussion on the selected families of vascular
plants, without sacrificing the discussion on basic principles. The families included for
discussion are largely those which have wide representation, as also those that are less
iv Plant Systematics
known but significant in evaluating the phylogeny of angiosperms. The discussion of the
families also has a considerable focus on their phylogenetic relationships, as evidenced by
recent cladistic studies, with liberal citation of molecular data. Several additional families
have been included for detailed discussion in the present volume.
Recent internet revolution has greatly helped in propagating taxonomic information,
with numerous searchable databases, online programs for identification and data analysis
available for ready reference. The information concerning these has been included at
appropriate places in various chapters for easy utilization. In light of this, the separate
chapter on web has been omitted. The outputs of computer programs, especially used in
molecular studies and construction of phylogenetic trees has been included based on actual
or hypothetical data. This will acquaint readers with the handling of raw data and working
of computer programs.
The author has attempted to strike a balance between classical fundamental
information and the recent developments in plant systematics. Special attention has been
devoted to the information on botanical nomenclature, identification and phylogeny of
angiosperms with numerous relevant examples and detailed explanation of the important
nomenclatural problems. An attempt has been made to present a continuity between orthodox
and contemporary identification methods by working on a common example. The information
on methods of identification using computers has been further enhanced to help better on-
line identification.
For providing me inspiration for this book, I am indebted to my undergraduate students,
who helped me to improve the material through frequent interactions. I am also indebted
to my wife Mrs. K.G. Singh for constant support and bearing with my overindulgence with
this book. I also wish to acknowledge the help rendered by my son Kanwarpreet Singh at
various stages.
I wish to record thanks to all the colleagues whose inputs have helped me to improve
the information presented here. I also wish to place on record sincere thanks to Dr. Jef
Veldkamp for valuable information on nomenclature, Dr. Gertrud Dahlgren for photographs
and literature, Dr. P.F. Stevens for literature on APG II and trees from his APweb, Dr.
Robert Thorne for making available his 2007 classification, Dr. James Reveal for his help
on nomenclatural problems, Dr. D.L. Dilcher for his photograph, Dr. Julie Barcelona and
Harry Wiriadinata for photographs of Rafflesia, the authorities of New York Botanical Garden,
Missouri Botanical Garden, USA, Royal Botanic Gardens Kew and University of California,
Santa Cruz, for photographs used in the book.

New Delhi Gurcharan Singh
November 2009
Chapter 6
Contents
Preface iii
1. PLANTS, TAXONOMY AND SYSTEMATICS 1–14
Plants and Kingdoms of Life 1
Two Kingdom System 1
Two Empires Three Kingdoms 2
Five Kingdom System 2
Six or Seven Kingdoms? 2
The Plant Kingdom 6
Taxonomy and Systematics 7
Basic Components (Principles) of Systematics 8
Aims of Systematics 11
Advancement Levels in Systematics 12
2. BOTANICAL NOMENCLATURE 15–45
Need for Scientific names 15
Why Latin? 16
Development of Botanical Code 16
Contents of Botanical Code 17
Preamble 17
Principles of ICBN 18
Names of Taxa 18
The Type Method 23
Author Citation 25
Publication of Names 26
Rejection of Names 28
Principle of Priority 30
Names of Hybrids 34
Names of Cultivated Plants 35
vi Plant Systematics
Unified Biological Nomenclature 35
Draft BioCode 36
PhyloCode 38
3. HIERARCHICAL CLASSIFICATION 46–55
Taxonomic groups, categories and ranks 46
Utilization of categories 48
Species concept 49
Infraspecific ranks 53
Genus 54
Family 54
4. DESCRIPTIVE TERMINOLOGY 56–90
Habit and life span 56
Habitat 57
Roots 57
Stems 58
Leaves 61
Leaf arrangement 62
Leaf duration 63
Leaf incision 63
Stipules 65
Leaf shape 65
Leaf margin 66
Leaf base 66
Leaf apex 67
Leaf surface 68
Venation 69
Inflorescence 69
Racemose types 69
Cymose types 70
Specialized types 71
Flower 71
Calyx 73
Corolla 74
Perianth 74
Androecium 74
Gynoecium 77

Fruit 79
Simple fruits 80
Aggregate fruits 82
Multiple fruits 83
Floral formula 83
Floral diagram 83
5. PROCESS OF IDENTIFICATION 91–127
Specimen preparation 91
Fieldwork 91
Equipment 92
Collection 93
Pressing 93
Contents vii
Handling special groups 94
Drying 94
Herbarium methods 95
Botanical gardens 95
Herbaria 101
Pest Control 105
Virtual herbarium 106
Identification methods 108
Taxonomic literature 108
Taxonomic keys 113
Cmputers in identification 120
Interactive keys Id. 121
6.VARIATION, BIOSYSTEMATICS, POPULATION GENETICS
AND EVOLUTION 128–148
Types of variation 128
Developmental variation 129
Environmental variation 129
Genetic variation 129
Variance analysis 129
Reproductive systems 131
Outbreeding 131
Inbreeding 135
Apomixis 135
Population genetics 135
Allele frequencies 136
Mating systems 136
Hardy-Weinberg law 136
Evolution 139
Mutation 140
Migration 140
Random genetic drift 140
Natural selection 141
Molecular evolution 143
Neutral theory of evolution 143
Speciation 144
7. TAXONOMIC EVIDENCE 149–209
Morphology 149
Habit 149
Underground parts 150
Leaves 150
Flowers 150
Fruits 150
Anatomy 150
Wood anatomy 150
Trichomes 151
Epidermal features 153
Leaf anatomy 153
Floral anatomy 153
viii Plant Systematics
Embryology 154
Families marked out by distinct embryological features 154
Specific examples of the role of embryological data 155
Palynology 156
Pollen aggregation 156
Pollen wall 157
Pollen aperture 157
Micromorphology and Ultrastructure 159
Micromorphology 159
Ultrastructure 161
Chromosomes 164
Chromosomal number 164
Chromosomal structure 167
Chromosomal behaviour 168
Chemotaxonomy 168
Primary metabolites 169
Secondary metabolites 169
Non-semantide Macromolecules 178
Proteins 178
Molecular systematics 184
Molecular evolution 184
Location of molecular data 186
Molecular techniques 187
DNA polymorphism 199
Examples of molecular studies 204
Gene trees 209
8. DEVELOPING CLASSIFICATIONS 210–264
Phenetic methods 210
Principles of taxometrics 211
Cladistic methods 212
Phylogenetic terms 213
Phylogenetic diagrams 221
Phylogeny and classification 225
Phylogenetic data analysis 229
Taxa-Operational Units 229
Characters 229
Measure of similarity 234
Construction of trees 237
The Consensus tree 251
Automated trees 258
Gene trees and species trees 262
Developing classification 263
9. PHYLOGENY OF ANGIOSPERMS 265–296
Origin of Angiosperms 265
What are Angiosperms? 265
What is the age of Angiosperms? 266
What is the place of their origin? 268
Are angiosperms monophyletic or polyphyletic? 270
Contents ix
What are the possible ancestors? 270
Origin of monocotyledons 280
Basal living angiosperms 280
Casuarinaceae 281
Magnoliids 281
Paleoherbs 282
Evolutionary trends 285
Coevolution with animals 285
Basic evolutionary trends 286
Xylem evolution 287
Stamen evolution 289
Pollen grain evolution 292
Carpel evolution 292
Evolution of inferior ovary 296
10. SYSTEMS OF CLASSIFICATION 297–358
Classifications based on gross morphology 297
Preliterate mankind 297
Early literate civilisations 297
Medieval Botany 299
Renaissance 300
Sexual System 302
Carolus Linnaeus 303
Natural Systems 306
Michel Adanson 306
Jean B. P. Lamarck 306
de Jussieu family 306
de Candolle family 307
Robert Brown 308
George Bentham and Sir J. D. Hooker 308
Phylogenetic Systems 312
Transitional Systems 312
Intentional phylogenetic systems 316
Modern phylogenetic systems 324
11. FAMILIES OF PTERIDOPHYTES 359–383
Lycopodiophtes
Lycopodiaceae 362

Selaginellaceae 363
Isoetaceae 365
Psilopsida
Ophioglossaceae 366
Psilotaceae 368
Equisetopsida
Equisetaceae 370
Pteropsida
Osmundaceae 371
Marsileaceae 373
Salviniaceae 374
Cyatheaceae 376
Pteridaceae 377
x Plant Systematics
Aspleniaceae 379
Dryopteridaceae 380
Polypodiaceae 382
12. FAMILIES OF GYMNOSPERMS 384–406
Cycadales
Cycadaceae 386
Zamiaceae 387
Ginkgoales
Ginkgoaceae 389
Coniferales
Pinaceae 391
Cupressaceae 393
Podocarpaceae 395
Araucariaceae 396
Taxaceae 398
Gnetales
Ephedraceae 399
Gnetaceae 401
13. MAJOR FAMILIES OF ANGIOSPERMS 407–678
Angiosperms roll of honour 408
Chloranthidae 409
Amborellaceae 409
Chloranthaceae 411
Austrobaileyaceae 413
Winteraceae
Illiciaceae 415
Cabombaceae 417
Nymphaeaceae 419
Ceratophyllaceae 421
Magnoliidae 423
Magnoliaceae 423
Degeneriaceae 425
Annonaceae 427
Calycanthaceae 429
Lauraceae 431
Winteraceae 433

Saururaceae 435
Piperaceae 437
Alismatidae 439
Acoraceae 439
Araceae 441
Butomaceae 443
Alismataceae 445
Hydrocharitaceae 447
Potamogetonaceae 449
Liliidae 451
Pandanaceae 451
Dioscoreaceae 453
Smilacaceae 455