# Plant succession; an analysis of the development of vegetation

Index Page
​Plant Succession An Analysis of the Development of Vegetation
By Frederic E. Clements
Professor of Botany in the University of Minnesota
Published by the Carnegie Institution of Washington
Washington, 1916
​Preface iii List of illustrations X

## Concept and Causes of SuccessionConcept and Causes of Succession

       The formation an organism 3
       Universal occurrence of succession 3
       Viewpoints of succession 3
       Succession and sere 4
       Sere and cosere 4
       Processes in succession 4

### Causes of SuccessionCauses of Succession

       Relation of causes 5
       Kinds of causes 5
       Proximate and remote causes 5

### Essential Nature of SuccessionEssential Nature of Succession

       Developmental aspect 6
       Functional aspect 7

## General Historical SummaryGeneral Historical Summary
### Early InvestigationsEarly Investigations

       King, 1685 8
       Degner, 1729 9
       Buffon, 1742 9
       Biberg, 1749 10
       Anderson, 1794 10
       De Luc, 1806 10
       Rennie, 1810 12
       Dureau, 1825 13
       Steenstrup, 1842 14
       Reissek, 1856 16
       Vaupell, 1857 17
       von Post, 1861 17
       Gremblich, 1876 18
       Müller, 1878-1887 19
       Other investigations 19

### Recent InvestigationsRecent Investigations

       Blytt, 1876 21
       Hult, 1885-1887 22
       Warming, 1891 23
       MacMillan, 1894-1896 24
       Warming, 1895 25
       Graebner, 1895 25
       Pound and Clements, 1898-1900 26
       Schimper, 1898 26
       Cowles, 1899 27
       Cowles, 1901 28
       Clements, 1902-1904 29
       Clements, 1904 29
       Friih and Schroter, 1904 30
       Clements, 1905-1907 30
       Moss, 1907-1910 30
       Clements, 1910 31
       Cowles, 1911 31
       Shantz, 1911 31
       Tansley, 1911 31
       MacDougal, 1914 32

## Initial CausesInitial Causes

       Significance of bare areas 33
       Modifications of development 33
       Processes as causes 34
       Change of conditions 34
       Fundamental nature of water-content 34
       Kinds of initial causes 35
       Physiography 36

### Topographic CausesTopographic Causes

       Topographic processes 36
       Kinds of processes 37
       Base-leveling 38

### ErosionErosion

       Nature 39
       Agents of erosion 39
       Rate and degree of erosion 40
       Fragmentary and superficial erosion 40
       Bare areas due to water erosion 41
       Bare areas due to wind erosion 41
       Bare areas due to gravity 41
       Bare areas due to ice action 41

### DepositDeposit

       Significance 42
       Agents of deposit 42
       Manner of deposit 43
       Rate and depth of deposit 43
       Place of deposit 44
       Distance of transport 44
       Fragmentary and local deposit 45
       Sterility of deposits 45
       Bare areas due to deposit by moving water 45
       Bare areas due to waves and tides 46
       Composition and water-content of alluvial deposits 46
       Bare areas due to deposit by ground waters 46
       Bare areas due to deposit by wind 47
       Deposit by ice and snow 47
       Bare areas due to deposit by glaciers 47
       Bare areas due to deposit by ice and snow 48
       Bare areas due to deposit by gravity 48
       Bare areas due to volcanic deposits 49
       Ponding and draining 49
       Kinds of lakes and ponds 50
       Life-history of a lake 50
       Drainage 51

### Elevation and SubsidenceElevation and Subsidence

       Elevation and subsidence 52
       New areas due to elevation 52
       Subsidence 52
       Earthquakes 54
       Similarity of topographic processes 54

### Edaphic CausesEdaphic Causes

       Nature 55 ​

### Climatic CausesClimatic Causes

       Róle 55
       Bare areas due to climatic factors directly 55
       Bare areas due to drouth 56
       Bare areas due to wind 56
       Bare areas due to snow, hail, and frost 56
       Bare areas due to lightning 57
       Bare areas due indirectly to climatic factors 57
       Sudden changes of climate 57

### Biotic CausesBiotic Causes

       General relations 58
       Action and effect 58
       Bare areas due to destruction of vegetation alone 59
       Bare areas with dry or drier soils 59
       Bare areas with wet soils or water 60

### Primary and Secondary AreasPrimary and Secondary Areas

       Distinction 60
       Sterility of primary and secondary areas 60
       Denudation 61
       Methods of denudation 61
       Depth of removal or deposit 61
       Rate and extent of removal 62

## Ecesic CausesEcesic Causes

       Nature 63

### AggregationAggregation

       Concept and róle 63
       Effects of simple aggregation 63
       Relation to denuded areas 64
       Interaction of aggregation and migration 64

### MigrationMigration

       Concept 64
       Mobility 64
       Seed-production 65
       Influence of the organ used 65
       Influence of the migration contrivance 66
       Role of migration agents 67
       Destructive action of agents 67
       Direction of migration 67

### EcesisEcesis

       Nature and róle 68
       Germination 69
       Fate of seedling 70
       Growth 71
       Reproduction 71
       Ecesis in bare areas 71

### CompetitionCompetition

       Nature 72
       Competition and dominance 72
       Competition in air and in soil 73
       Róle of competition in succession 73

### InvasionInvasion

       Nature and róle 75
       Kinds of invasion 75
       Manner of invasion 76
       Barriers 77
       Biological barriers 77
       Changes in barriers 78

## ReactionsReactions

       Concept and nature 79
       Róle in succession 80
       Previous analyses of reaction 80
       Kinds of reactions 81

### Soil FormationSoil Formation

       Manner 81
       Reaction by accumulating plant bodies or parts 81
       Reaction by accumulating plant concretions 83
       Reaction by producing weathering 83
       Reaction upon wind-borne material 84
       Reaction upon water-borne detritus 85
       Reaction upon slipping sand and gravel 86

### Soil StructureSoil Structure

       Reaction by adding humus 86
       Reaction by compacting the soil 87
       Reaction by preventing weathering or erosion 88

### Water-ContentWater-Content

       Reaction by increasing water-content 88
       Reaction by decreasing water-content 89

### Nutrients and SolutesNutrients and Solutes

       Reaction by adding nutrients or foodstuffs 89
       Reaction by decreasing nutrients 89
       Reaction by producing acids 90
       Reaction by producing toxins 90

### Soil OrganismsSoil Organisms

       Reaction by means of parasites 91
       Reaction by means of saprophytes 92

### Air ReactionsAir Reactions

       Reaction upon light 92
       Reaction upon humidity, temperature, and wind 94
       Reaction upon local climate 94
       Reaction upon aërial organisms 95
       Correlation of reactions 96
       Quantitative study of reactions 96

## Stabilization and ClimaxStabilization and Climax

       Stabilization 98
       Causes of stabilization 98
       Relation to the climax 98
       Degree of stabilization 99

### Life-History StagesLife-History Stages

       Nature
       Kinds of stages 100
       Role of life forms 100
       Reasons why plants disappear 100
       Reasons why plants appear at certain stages 100
       Reasons why plants appear before their proper time 103
       Initial stages 103
       Medial stages 105 ​

### The ClimaxThe Climax

       Concept 105
       Nature 106
       Relation to succession 106
       Kinds of climaxes 107
       Subclimaxes 107
       Potential climaxes 108
       Changes of climate 109
       Preclimax and postclimax 109
       Changes of climax 110

## Structures and Units of VegetationStructures and Units of Vegetation
### Development and StructureDevelopment and Structure

       Relation 111
       Kinds of structure 111
       Zonation 111
       Relations of climax zones 113
       Significance of alternation 115
       Developmental relation of layers 115
       Relation of seasonal aspects 116

### The Units of VegetationThe Units of Vegetation
#### Historical Summary:Historical Summary:

       The formation concept 116
       Grisebach's concept of the formation 116
       Drude's concept 117
       Clements's concept 118
       Moss's concept 118
       Schröter's concept 119
       Gradmann's concept 119
       Warming's concept 120
       Negri's concept 122
       Correlation of divergent views 122
       Significance of development 123
       Earlier suggestions of developmental view 123

#### The Formation:The Formation:

       Developmental concept of the formation 124
       Analysis of the formation 125
       Formation units 125
       Formation 126
       Names of formations 127

#### Climax Units:Climax Units:

       Association 128
       Consociation 129
       Society 130
       Bases 131
       Kinds of societies 132
       Aspect societies 132
       Layer societies 133
       Cryptogamic societies 133
       Terminology 133
       Changes of rank or dominance 134
       Clan 134

#### Seral Units:Seral Units:

       Nature and significance 135
       Associes 136
       Consocies 137
       Socies 138
       Colony 138
       Family 139
       Summary of units 139
       Mixed communities 139
       Nomenclature of units 140
       Formation groups 142
       Bases 143
       Developmental groups 143

## Direction of DevelopmentDirection of Development

       Development always progressive 145
       Nature of regression 145
       Course of development 146
       Regression and retrogression 146
       Nilsson's view 146
       Cowles's view 147
       Cajander's view 149
       Sernander's view 149
       Moss's view 150
       Hole's view 154
       Conversion of forest 155
       Status of forest in Britain 156
       Artificial conversion 157
       Graebner's studies: Conversion of forest to heath 157
       Conversion of forest into moor 159
       Causes of conversion 161
       Possibility of backward development 162
       Degeneration 163
       Regeneration 163
       Correlation of progressive developments 164
       Convergence 165
       Normal movement 166
       Divergence 167

## Classification of SeresClassification of Seres

       Historical 168
       Clements's system 168
       Normal and anomalous succession 169
       Primary and secondary succession 169
       Cowles's system 171
       Possible bases of classification 175
       Developmental basis of classification 175
       Initial areas and causes 176
       Relative importance of bases 177
       The climax as a basis 177
       Recognition of climax areas 177
       Climaxes of North American vegetation 179
       Subclimaxes 180
       Relationship of climaxes 180
       Names of climaxes 181
       Priseres and subseres 182
       Hydroseres and xeroseres 182
       Phylogenetic system 183

## Climax Formations of North AmericaClimax Formations of North America

       Classification 184

### The Deciduous Forest ClimaxThe Deciduous Forest Climax

       Prisere 184
             Hydrosere 184
             Xerosere 203
       Subsere 212

### The Prairie-Plains ClimaxThe Prairie-Plains Climax

       Prisere 214
             Xerosere 214
             Hydrosere 220
       Subsere 221

### The Cordilleran ClimaxesThe Cordilleran Climaxes

       Priseres 225
       Subseres 226 ​
       Desert Climaxes
       Southeastern Climaxes

## Succession in EurasiaSuccession in Eurasia

       Scandinavia
       Britain
       Middle Europe
       Russia
       Mediterranean region
       Tropics and subtropics

## Past Climates and ClimaxesPast Climates and Climaxes
### InterpretationInterpretation

       Interpretation of past vegetations 279
       Methods of interpretation 280

### Vegetation ErasVegetation Eras

       Criteria 283
       Evidence from the method of sequence 286
       Division into eras 288
       Saporta's table of eras and periods 288
       Relation of vegetation era and eosere 289
       Comparison of geologic and vegetation eras 289

### The Plant RecordThe Plant Record

       Nature 290
       Strates and stases 291
       The strate 292
       Kinds of strates 293
       The stase 293
       Water stases 294
       Kinds of stases 295
       Relations of strate and stase 295
       Relation to stratigraphic units 296
       Relation to succession 297

### Deformation and GradationDeformation and Gradation

       Processes 297
       Nature of deformation 298
       Great periodic movements of deformation 298
       Sequences of deformation 299
       Permian deformation 300
       The circulation sequence 300
       The gradation sequence 300
       The climatic sequence 301
       The deformation cycle 302
       The series of deformation cycles 304
       Deformation and unconformity 305
       Correlation of climates 306
       Effects upon vegetation 307
       The cycle of erosion 307
       Relation to vegetation 309

### Past ClimatesPast Climates

       Interpretation 312
       Evidence of past climates and changes 313
       Geologic evidence 313
       Salt and gypsum 314
       Red beds 314
       Terraces 315
       Fluctuations of lake levels 316

### Botanic EvidenceBotanic Evidence

       Plant fossils 317
       Successional evidences 317
       Evidence from stases 318
       Evidence from annual rings 318

### Zooic EvidenceZooic Evidence

       Animal fossils 319
       Culture relicts 319

### Causes of Climatic ChangesCauses of Climatic Changes

       Kinds of causes 320
       The deformational hypothesis 321
       The volcanic hypothesis 322
       The solar hypothesis 325
       Kullmer's law of the shift of the storm-track 326
       The cyclonic solar hypothesis 327
       Relative value of causes 327
       Coincidence of causes 328

### Climatic CyclesClimatic Cycles

       Relation of changes and cycles 329
       Kinds of cycles 330
       Arotowski's cycle of 2½ years 330
       The 11-year sun-spot cycle 332
       Sun-spot maxima and minima from 1750 to 1913 333
       Table of maximum and minimum years, 1610 to 1913 334
       Nature of sun-spots 334
       Effects of sun-spots upon climate 334
       Douglass's 21-year cycle 336
       Brückner's 35-year cycle 336
       Major sun-spot cycles 337
       Volcanic cycles 338
       Deformational cycles 339

### The Series of Climax CyclesThe Series of Climax Cycles

       Correlation of cycles 339
       Table of climatic cycles 340
       Spatial differentiation of climates 341

### Correlation of Climatic Cycles and SuccessionCorrelation of Climatic Cycles and Succession

       General relations 342
       Responses of vegetation 342
       Relation to the different climatic cycles 343

## Past Succession: The CeneoserePast Succession: The Ceneosere
### General PrinciplesGeneral Principles

       Phylogeny and ontogeny 344
       Recapitulation 345
       Geosere and eosere 346
       Clisere 347
       Cosere 348
       Correlations of seres 349
       Processes and principles of past succession 349 ​

### The Cenophyiic EraThe Cenophyiic Era

       The Flora:
             Nature and origin 351
             Relation to Mesophytic and Paleophytic floras 351
             Inferences from distribution 352
             Inferences from phylogeny 353
       The Life-forms:
             The record 354
             Methods of inference 354
             Dominants 355
             Structure of the vegetation 356
       Climates and Habitats:
             Relation of habitat to climate 357
             Deformational and climatic cycles 359
             Periods and epochs 359
       The Ceneosere:
             The Cretaceous period 360
             Life-forms and dominants 361
             Cretaceous seres and coseres 363
       The Cretaceous-Eocene clisere:
             The deformation cycle 364
             Vegetation zones 365
             Dominants of the Eocene 366
       The Oligocene-Miocene clisere
             The deformation cycle 365
             Coseres and cliseres 366
       The Pleistocene cliseres and coseres
             The deformation cycle 368
             The causes of glaciation 368
             Glaclal-interglacial cycles 371
             The Pleistocene flora 371
             Pliocene climax zones 372
             The Jerseyan-Aftonian clisere 372
             Seres and coseres 376
             Postglacial succession 377
       The Peat Clistase 378

## Mebeosere and PaleosereMebeosere and Paleosere
### The Mesophytic EraThe Mesophytic Era

       The flora 404
       Composition of the flora 404
       Life-forms and dominants 405
       Structure of the vegetation 406
       Deformation and climatic cycles 406
       Climate of the periods 408

### The MeseosereThe Meseosere

       Permian succession 409
       Triassic succession 411
       Jurassic succession 412
       The Cycadean climax 413
       Comanchean succession 414

### The Paleophytic EraThe Paleophytic Era

       The flora 417
       Life-forms and dominants 419
       Structure of the vegetation 419
       Deformational cycles and climates 419

### The PaleosereThe Paleosere

       The sere 420
       The cosere 421
       The clisere 422

## The Investigation of SuccessionThe Investigation of Succession

       Primary methods 423
       Special methods 424

### The Quadrat MethodThe Quadrat Method

       Concept and significance 424
       Kinds of quadrats 425
       List quadrat 425
       Chart quadrat 426
       Permanent quadrat 427
       Denuded quadrat 429
       Quadrat series and sequences 429
       Various quadrats 430
       The transect 430
       The bisect 432
       The migration circle 433

### Methods of MappingMethods of Mapping

       Methods 435
       Community charts and ecotone maps 435
       Survey maps 436
       Climax maps 436

### Instrumental MethodsInstrumental Methods

       General considerations 437
       Measurement of reactions 438
       Measurement of water reactions 438
       Measurement of light reactions 439

### Growth MethodsGrowth Methods

       Ring counts 440
       Burn-scars 441

## BibliographyBibliography

==Index==