cover

The Biology of Giant Kelp Beds (Macrocystis) in California

Ed.: Wheeler J. North

1970. 600 pages, 166 figures, 116 tables, 17x24cm, 1500 g
Language: English

(Nova Hedwigia, Beihefte, Beih. 32)

ISBN 978-3-768-25432-8, paperback, price: 103.00 €

out of print

BibTeX file

Contents

Synopsis top ↑

This book is the last of three large publications resulting from kelp studies conducted recently at the University of California's Institute of Marine Resources (IMR). The two preceding volumes are ”An inves— tigation of the effects of discharged wastes on kelp” and ”The utilization of kelp bed resources.” Both of these works, which emphasize practical aspects of the kelp environment, are discussed briefly in the next section below. The present publication focuses on the general biology of giant kelp and the organisms associated with it.
Each author is an active researcher in the topic he discusses. Rather than review literature extensively, emphasis has been placed on presenting new material. Nonetheless the individuals have tried to review their fields adequately, though briefly. Unfortunately such independently written chapters may yield a piecemeal presentation and lack coherency. As a remedy, our introductory chapter will present a more unified and general view of the present state of knowledge in kelp biology. We have also included certain subjects outside the scope of pure biology since they will add significantly to the completeness of our presentation (economics of kelp resources, other IMR kelp publications).
Although our discussions are as complete as possible, the reader will perceive that in some fields much information is sketchy or unavailable. Species lists, measures of importance, and identification keys need to be extended for many important but poorly studied groups of the kelp bed assemblage (tunicates, canopy crustaceans, sponges, hydroids, etc.). Distribution of organisms as a function of geographical range and of depth should be studied. The ecology of the Macrocystis gametophyte is very poorly understood, as is the entire field of microbiology of kelp beds. These and many other conspicuous problems point up how little we still know about our oceanic forests.

Table of Contents top ↑

Frontispiece XV
1. Introduction and background. W. I. North 1
Scope and organization 1
Other publications of the IMR Kelp Program 2
The utilization of kelp bed resources 2
The effects of discharged wastes on kelp 4
Early interests 5
The Macrocystis plant 12
Factors limiting distribution 12
Local limitations 12
Worldwide limitations 14
Systematics 16
Life history and development 17
Microscopic stages 17
Early sporophyte 18
Young and mature sporophytes 20
The kelp frond 24
Development 24
Cross morphology of the mature frond 29
Morphological variation 29
Physiological features 30
Fruiting blades 35
Lifespan of the fronds 35
Lifespan of the plant 36
Ecology of kelp beds 37
The Macrocystis niche 37
Ecological successions 42
Regular changes 42
Non-regular changes 42
Lack of changes 44
Biomass and concentrations of Macrocystis 45
Biomass — methods and results 45
Frond concentrations 46
Plant concentrations 46
Associated vegetation 46
Modifications due to Macrocystis 46-
Seaweed species in southern California kelp beds 46
Central, southern, and Baja California kelp beds 48
Associated fauna 52
Historical aspects 52
Animal species in southern California kelp beds 54
Sessile filter feeders 64
Grazing animals 66
Carnivores and Scavengers 66
Nourishment of animals outside of kelp beds 66
Kelp as substrate and shelter 67
Resource aspects of kelp beds 68
Origins and history of the California kelp industry 68
Plant resources 72
Associated algae 72
Composition and current uses of Macrocystis 72
Algin 72
Animal resources 78
Marine waste disposal 83
Economic values of the resources 85
Kelp 86
Commercial fishes and shellfish 88
Sportfishes 89
Waste disposal value of the ocean 93
Value comparisons 96
Acknowledgements 96
2. The internal structure of Mncrocystis. B. C. Parker 99
Introduction 99
The apical region of the mature frond 99
The mature stipe 103
The holdfast 116
The blades 119
Acknowledgements 121
3.Growth of individual fronds. W. I. North 123
Introduction 123
Growth in Macrocystis 124
Methods 125
Measurements of length 126
Measurements of weight 128
Measurements of laminar area 129
Results 129
Historical method 130
Synoptic method 130
Comparison of the Two Methods 131
Weight and laminar increments 138
Equations describing average elongation 140
Variability of standard growth rates 142
Total variability 142
Individual variability 145
Long term changes 147
Effect of temperature 149
Effect of depth 152
Effect of bed density 154
Effect of plant age 156
Geographical effects 160
Effects of season 163
Summary 165
Acknowledgements 168
4. Photosynthesis and general development in Macrocystis. K. A. Clendenning 169
Photosynthesis 169
Introduction 169
Decentralization of photosynthesis and growth in kelp 170
Photosynthetic saturation of bottom tissues 172
The kelp blade 172
Ontogenetic changes 175
Effect of high light intensities 175
Effect of short exposures to high temperatures 176
Intertidal kelp from Turtle Bay, Mexico 177
Action of antibiotics 177
Effect of blade damage and of cutting 177
Daily time course of photosynthesis 178
Seasonal changes 178
Development of young kelp fronds under dense surface canopies 183
Methods and results 184
Characteristics of first generation Macrocystis fronds 185
Regeneration and function of the apical ”scimitar” blade 187
Natural transport of Macrocystis 187
lntercalary growth in giant kelp fronds 188
Mechanism of surface canopy formation 188
5. Studies of translocation in Macrocystis. B. C. Parker 191
Introduction 191
Current research with C“ and fluorescein dye 193
Future research on translocation 193
6. Gross composition of kelp. K. A. Clendenning 197
Blade area, total solids, and organic matter of developing kelp fronds 197
Measurement of water and solids of the various organs in Macrocystis 199
Evaporation rates 200
Solids content of young blades 202
Solids content of mature blades 203
Solids content of kelp powder 203
Discussion 204
Ash content of kelp 204
Solids, ash, and organic matter in holdfasts and stipes 206
7. The species of Macrocystis. M. Neushul 211
Introduction 211
The species of Macrocystis 211
Species distribution 213
Macrocystis of South America 213
Macrocystis of North America 217
Discussion and conclusions 221
Acknowledgements 222
8. Macrocystis and related kelps. M. Neushul 223
9. Physical aspects of the nearshore environment. I. C. Quast 229
Introduction 229
Acknowledgements 229
Thekelpenvironment 231
Temperature 233
Wave action 235
Visibility 235
Discussion 238
Summary 240
10. Submarine illumination in Macrocystis beds. M. Neushul 241
Introduction 241
Measurement of submarine illumination 241
Results 245
Discussion and conclusions 251
Acknowledgements 254
11. Phytoplankton as light absorbing agents in the La Jolla kelp bed.
K. A. Clendenning 255
12. Organic productivity in kelp area. K. A. Clendenning 239
Organic matter synthesis by giant kelp beds 259
Kelp vs. phytoplankton productivity 262
13. The kelp community of seaweeds. M. Neushul 265
14. Drift seaweeds on San Diego County beaches. C. E. ZoBell 269
Introduction 269
Field observations 271
Stations occupied and data collected 271
Methods of estimating quantities of drift seaweeds 277
Kinds of seaweeds on beaches 282
Quantities of seaweeds on beaches 287
Seasonal abundance of beached seaweeds 291
Effects of water movements 291
Effect of beach conditions on abundance of beached seaweeds 295
Factors affecting occurrence of seaweeds in surf 298
Effects of kelp harvesting 301
Condition of beached seaweeds 307
Fate of drift seaweeds 309
Drift seaweeds on beaches elsewhere 311
Summary and conclusions 313
Acknowledgements 314
15. Microbiological aspects of the kelp bed environment. H. L. Scotten 315
General Findings 315
Bacterical characteristics 315
Kelp antibiotic activity 316
Temparature grouping 316
Kelp microflora near sewer outfalls 317
Studies of black rot 317
16. Kelp surfaces and associated invertebrates. B. L. Wing and K. A. Clendenning
319
Macrocystis surfaces throughout the water column 319
Membranipora encrustations on kelp 319
General description of the colonies 323
Feeding 324
Weight 324
Taxonomy 325
Photosynthesis and respiration in heavily encrusted kelp blades 320
Motile invertebrates of kelp fronds 327
Methods 327
Results 332
Discussion 339
Kelp blade perforations and attached foraminifera (assisted by R. R. Lankford)
340
17. Small herbivorous invertebrates of canopy and holdfast. L. G. Jones 343
Introduction 343
The kelp isopod, Idothea (Pentidothea) resecata (Stimpson) 344
Natural history 344
Quantitative feeding studies 347
Tropic relationships 350
Food preference studies 351
Temperature effects 351
Other Idothea species 353
A holdfast relative 353
A surfgrass relative 353
The pillbug Paracerceis cordata (Richardson) 353
The kelp curler, Ampitoe humeralis Stimpson 354
The kelp snail Mitrella carinata Hinds 354
The chink shell Lacuna unifas'ciata Carpenter 357
The gribble, Limnoria 358
Taxonomy 358
Natural history 359
Field observations of holdfast burrowing 362
Quantitative feeding studies 363
Preference experiments 365
Seasonal migrations 366
Summary 366
18. Mysids of the southern kelp region. W. D. Clarke 369
Introduction 369
Spatialdistribution 371
Discussion of individual species 373
Siriella pacifica 373
Acanthomysis sculpta 376
Acanthomysis columbiae 376
Acanthomysis macropsis 377
Acanthomysis costata 377
Acanthomysis 5p. 378
Neomysis rayi 378
Neomysis kadiakensis 378
Proneomysis Wailesi 378
Metamysidopsis elongata 378
Mysidopsis californica 378
Mysidopsis spp. 378
Heteromysis odontops 379
Bionomics of mysids 379
Mysids as food of fish 379
19. Species structure of the holdfast community. R. J. Ghelardi 381
Introduction 381
Locality, habitat description, and methods 381
Thestudyarea 381
I-Ioldfast description and sampling 383
Identification 386
Results 387
Thesub-habitats 387
Principal faunal groups 392
Gammarid amphipods 392
Frequentgammarids 392
Total species 396
Polychaetes 396
Frequent polychaetes 396
Less frequent polychaetes 399
Total species 399
Isopods 400
Frequent isopods 400
Total species 401
Miscellaneous 402
Ophiuroidea 404
Caprellidea 404
Chelifera 404
Higher taxonomic categories 406
Total species 415
Discussion 416
Summary 420
Acknowledgements 420
20. Grazing activities of benthic invertebrates in kelp beds. D. L. Leighton 421
Introduction 421
Studies of grazing in the field 425
Feeding habits of the chief grazer species 434
Orientation toward scent of Macrocystis 434
Food preferences 435
Feeding rates 438
Influence of temperature 438
Influence of animal size 439
Species comparisons 439
Nutritional values 441
Various Algae 441
Growth studies of sea urchins 443
Discussion 445
Grazing potentials of each species 446
Purple urchin 446
Giant red urchin 446
White urchin 447
Southern kelp crab 447
Common kelp crab 447
Smooth brown turban 447
Wavy top 447
Abalone 448
Sea hare 448
Macrocystis vs. other algae 448
Relations of temperature to grazing pressure 449
Grazing rates 450
Natural and artificial controls of grazing populations 451
Summary 452
21. Bryozoans of California and northern MexiCo kelp beds. R. M. Woolacoti
and W. I. North 455
Introduction 455
Bryozoan species from kelp areas 459
Widely distributed species 463
Localized species 467
Rare species 469
Non-categorized species 470
Species frequency 472
Frequency 60 to 80 percent 473
Frequency 40 to 60 percent 474
Frequency 20 to 40 percent 474
The important species 474
Summary 476 F
Acknowledgements 479
22. Fish fauna of the rocky inshore zone. I. C. Quast 481
Introduction 481
Methods 481
The rocky-inshore fauna 481
General ecology 490
The fishcommunity 492
Summary 498

23. Estimates of the populations and standing crop of kelp bed fishes. J. C. Quast 509
Introduction 509
Methods 510
The belt transect 510
The wall net 513
Estimation of the fish standing crop 516
Belt transect surveys 516
Evaluation of belt transect estimates 52-4
Quantitative collections 525
Combined estimates 532
Summary 536
Appendix A: cost of wall net materials 538
Appendix B: weight-length relationships of common kelp bed fishes 539
24. Observations on the food of kelp bed fishes. J. C. Quast 541
Introduction 541
Acknowledgements 541
Methods 542
Findings 542
California moray 542
California halibut 544
Kelp bass 544
Sand bass 544
Topsmelt 546
Pacific barracuda 547
Sargo 547
Salerna 548
White seabass 548
Black croaker 548
Ocean Whitefish 550
Kelp perch 550
Black perch 552
White seaperch 552
Rubberlip seaperch 553
Pileperch 553
Blacksmith 553
Garibaldi 554
Rock wrasse 555
Senorita 556
Sheephead 559
Opaleye 559
Halfmoon 562
Zebraperch 562
California scorpionfish 562
Kelp rockfish 563
Brown rockfish 563
Gopher rockfish 565
Vermilion rockfish 565
Blue rockfish 565
Grass rockfish 567
Olive rockfish 568
Treefish 569
Cabezon 569
Lingcod 570
Painted greenling 571
Slimy snailfish 572
Kelp pipefish 572
Island kelpfish 572
Spotted kelpfish 573
Striped kelpfish 574
Giant kelpfish 575
Reef finspot 575
Mussel blenny 575
Spotted cusk eel 576
Importance of the foods 576
References 581