The Ecology of Seashores - Chapter 1 docx

Earlywork on seashores concentrated on the problems of life in an environment characterized by steep gradients inphysical conditions, but in more recent years, the focus of research on t

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Seashores

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George A Knox, M.B.E., F.R.S.N.Z.

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Knox, G A.

The ecology of seashores / by George A Knox.

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Because of its accessibility, the intertidal zone has offeredexcellent opportunities to study the adaptations of indi-vidual organisms and populations to their environment,and the factors controlling community composition Earlywork on seashores concentrated on the problems of life

in an environment characterized by steep gradients inphysical conditions, but in more recent years, the focus

of research on the fascinating shore ecosystems has been

on understanding the processes controlling their tivity and dynamic functioning The emphasis has been

produc-on processes such as primary productiproduc-on, grazing, tion, competition, impact of disturbance, secondary pro-duction, detritus formation, decomposition, and the role

preda-of microorganisms

My own involvement in seashore research began when

I embarked on a M.Sc thesis in zoology on the ecology

of the rocky shores at Taylors Mistake, Banks Peninsula,with special reference to the serpulid polychaete Pomato-

I became involved in research on a local estuary, the Heathcote Estuary This research evolved into a compre-hensive interdisciplinary research program that has con-tinued until the present time It has involved a large num-ber of research associates, research assistants, andresearch students and culminated in two major reports(Knox and Kilner, 1973; Knox, 1992) A multiauthor bookbringing together the results of 50 years of research onthis estuarine ecosystem is in an advanced stage (Knoxand Robb, in preparation) Over the period of 1959 to

Avon-1983, I directed the activities of the Estuarine ResearchUnit, Zoology Department, University of Canterbury,Christchurch This unit carried out research on estuarineand coastal ecosystems throughout New Zealand and com-piled some 28 major reports The research aimed at under-standing the interaction of estuarine plants, microorgan-isms, animals, and man with each other and theirenvironment Such research, while contributing to basicecological principles, also provided information necessaryfor the management of New Zealand coastal ecosystems(Knox, 1983c) I was scientific coordinator of a multidis-ciplinary study of the Ahuriri Estuary, Hawke Bay (Knoxand Fenwick, 1978; Knox, 1979b), and coordinator of thebiological aspects of the Upper Waitemata Harbour Catch-ment Study, a comprehensive five-year interdisciplinarystudy of the land and water resources of the mangrove-fringed Upper Waitemata Harbour, Auckland (Knox,1983a; 1983b) A fellowship at the East-West Center,Honolulu, enabled me to undertake a study of coastal zone

resource development and conservation in Southeast Asiawith special reference to Indonesia (Knox and Miyabara,1983) This estuarine research culminated in 1983 withthe publication of a two-volume work on estuarine eco-systems (Knox, 1983a,b)

Over the years I have also been involved in research

on the intertidal ecology of rocky shores (Knox, 1963a;1968; 1969b; 1988; in preparation; Knox and Duncan,

in preparation) and sand beaches (Knox, 1969a) Fieldwork has been carried out on rocky shores throughoutNew Zealand, the Subantarctic Islands (Snares, Auck-land, Campbell, and Chatham Islands), Antarctica, thewest coast of Chile, and the southern coasts of Australia.Shores in Peru, Argentina, the eastern and western coasts

of Canada and the United States, England and Scotland,and tropical shores in Guam, the Palau Islands, Fiji, andHawaii have been briefly examined A book on NewZealand seashore ecology will be published shortly(Knox, in preparation) This work has also involved anexamination of the evolution and biogeography of theSouthern Hemisphere, especially the Pacific Ocean,intertidal and shallow water biotas (Knox, 1960; 1963a;1975; 1988)

The approach used in this book is somewhat differentfrom that used in many texts in marine ecology in that theemphasis is on ecological processes and the use of systemsanalysis in understanding such processes This book is anattempt to bring some order to some of the most complex

of ecosystem types, those of the seashores of the world.Wherever possible the energy circuit language of symbolsand diagrams developed by H T Odum has been used as

a basis of understanding (Odum, 1983)

The book is divided into seven chapters The firstprovides an introduction and sets the scene for the suc-ceeding chapters The second deals with zonation patterns

on hard shores, the basic causes of zonation, the biology

of the major divisions of the biota, the biology of somekey faunal components (mussels, limpets, and barnacles),and the flora and fauna of special habitats The thirdchapter covers the physicochemical environment of softshores, and the general ecology of the biota from produc-ers through microbes to macrofauna Two large sectionsdeal with microbial ecology and detritus and nutrientcycles in representative ecosystems The fourth chaptergives an account of ecological niches on the shore andthe establishment and maintenance of zonation patterns

as grazing, competition, predation, disturbance, and

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succession that determine the distribution, diversity, mass, and production of the various categories of con-sumers Energy budgets, patterns of energy flow, trophicstructure, and food webs are discussed in Chapter 6, while

var-ious ecosystem types on both hard and soft shores aredetailed Finally, the application of the relatively newtechnique of network analysis to gain greater insight intoecosystem processes and enable the comparison of differ-ent ecosystems is outlined

As will be seen by the reference list at the end of thisbook, there is a considerable volume of recent literature onseashore ecosystems, although the list contains only a frac-tion of the published work Extensive literature citationshave been included so that the book might serve as aresource for those engaged in research on and management

of the coastal zone Because of this growing volume of

literature, I have had to be selective in the material included.Thus, of necessity, I have concentrated on work publishedsince 1970, and in particular in the last decade Regretfully,except in a few instances, it has not been possible to developthe history of the concepts considered Examples have beencarefully chosen from the pool of published research toillustrate these concepts There are doubtless other exam-ples that could have been equally used, and I apologize toauthors whose work has not been included I have attempted

to include as wide a geographic range of examples as sible, and in particular have included Southern Hemisphereexamples that often do not appear in texts originating in theNorthern Hemisphere

pos-The book has been designed for use by upper levelundergraduate and graduate students and professionalsengaged in coastal zone research and management I hopethat they find it useful

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© 2001 by CRC Press LLC

Department of Zoology, University of Canterbury,Christchurch, New Zealand, from 1959 to 1976 He is nowprofessor-emeritus in zoology

Professor Knox was born in New Zealand andreceived his education at the University of Canterburywhere he was appointed a staff member in 1948 He hasbeen a visiting fellow at the East-West Center, Honolulu,and a visiting professor at the Department of Oceanogra-phy, Texas A&M University and the Department of Envi-ronmental Engineering, University of Florida, Gainesville

He has visited and worked in laboratories in the U.S.,Canada, Chile, Japan, Australia, Western Europe, theUSSR, and China

Professor Knox’s research has been wide ranging andincludes: (1) the systematics and distribution of polycha-eta with special reference to New Zealand and Antarctica;

(2) rocky shore intertidal ecology and biogeography; (3)the ecology and conservation of islands; (4) studies on thepelagic and benthic ecosystems beneath the sea ice inMcMurdo Sound, Antarctica; and (5) estuarine and coastalecology and management He established and directed theEstuarine Research Unit in the Department of Zoologyand the University of Canterbury Antarctic Research Unit

He has participated in many field expeditions, includingthe Chatham Islands 1954 Expedition (leader); the RoyalSociety of London Darwin Centennial Expedition tosouthern Chile (marine biologist and deputy leader); thir-

teen summer expeditions to McMurdo Sound, Antarctica;the establishment of the Snares Islands Research Pro-gramme (participated in three field expeditions); and par-ticipation in field expeditions to Campbell and AucklandIslands He has published over 100 scientific papers and

28 environmental reports, written five books, and editedand co-authored three other volumes

Professor Knox has received a number of awards andfellowships for his contributions to science, including Fel-low of the Royal Society of New Zealand (FRSNZ), 1963;Hutton Medal, Royal Society of New Zealand, 1978; Con-servation Trophy, New Zealand Antarctic Society; Mem-ber of the Most Excellent Order of the British Empire(MBE), 1985; New Zealand Marine Sciences SocietyAward for Outstanding Contribution to Marine Science inNew Zealand, 1985; and the New Zealand Association ofScientists’ Sir Ernest Marsden Medal for Service to Sci-ence, 1985

Throughout his career, professor Knox has been active

in international scientific organizations He has been amember of the Scientific Committee for Oceanic Research(SCOR) and the Special Committee for the InternationalBiological Programme (SCIBP) He has been a delegate

to the Scientific Committee for Antarctic Research (SCAR)since 1969, serving 4-year terms as secretary and president,and a member of the governing board of the InternationalAssociation for Ecology from 1965 to 1990, also serving4-year terms as secretary-general and president

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© 2001 by CRC Press LLC

Acknowledgments

I would like to express my indebtedness to the late ProfessorEdward Percival, who by his enthusiasm and teaching skillsstarted me on my career as a marine biologist, to ProfessorHoward Odum who inspired by interest in the energy anal-ysis approach to ecosystem modeling, and to my colleagues

in New Zealand and various parts of the world with whom

I have discussed many of the ideas in this book

My thanks are also due to all those who gave mepermission to reproduce original figures and tables Finally,

I am indebted to John Sulzycki and the staff of CRC Press,

in particular Amy Rodriguez and Pat Roberson, for theirpatience and support during the preparation of this volumeand for seeing the project through to the completion of such

a high quality product

George A KnoxNovember, 2000Christchurch, New Zealand

Chapter 1 The Environment

1.1 Introduction 1

1.2 Environmental Gradients and Stresses on the Shore 2

1.3 Salient Features of the Shore Environment 2

1.4 Patterns of Zonation on the Shore 16

Chapter 2 Hard Shores 2.1 Zonation Patterns on Hard Shores 20

2.2 Zonation Patterns on Representative Shores 24

2.3 The Causes of Zonation 36

2.4 Hard Shore Microalgae 51

2.5 Hard Shore Micro- and Meiofauna 53

2.6 Rocky Shore Lichens 54

2.7 Hard Shore Macroalgae 56

2.8 Key Faunal Components 58

2.9 Special Habitats 70

Chapter 3 Soft Shores 3.1 Soft Shores as a Habitat 87

3.2 The Physicochemical Environment 92

3.3 Soft Shore Types 96

3.4 Estuaries 98

3.5 Soft Shore Primary Producers 107

3.6 Soft Shore Fauna 153

3.7 Biological Modification of the Sediment 181

3.8 Microbial Ecology and Organic Detritus 188

3.9 Nutrient Cycling 209

3.10 Estuarine Shelf Interactions 228

Chapter 4 Adaptations to Shore Life 4.1 Introduction 238

4.2 Ecological Niches on the Shore 238

4.3 The Establishment of Zonation Patterns 251

4.4 The Maintenance of Zonation Patterns 263

Chapter 5 Control of Community Structure 5.1 Introduction 277

5.2 Hard Shores 277

5.3 Soft Shores 317

5.4 Synthesis of Factors Involved in Controlling Community Structure 344

Chapter 6 Energy Flow, Food Webs, and Material Cycling 6.1 Introduction 356

6.2 Food Sources 356

6.3 Energy Budgets for Individual Species 360

6.4 Optimal Foraging 374

6.5 Secondary Production 377

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6.6 P:B Ratios and Production Efficiency 380

6.7 Relative Contribution of Soft Shore Benthic Infauna to Secondary Production 381

6.8 Community Metabolism 385

6.9 Trophic Structure and Food Webs 389

6.10 Carbon Flow Models 406

6.11 Stable Isotopes and Food Web Analysis 412

6.12 Top-down and Bottom-up Control of Trophic Structure 421

Chapter 7 Ecosystem Models 7.1 Introduction 426

7.2 Hard Shores 427

7.3 Sand Beaches 428

7.4 Intertidal Mudflat in the Lynher Estuary, Cornwall, U.K 433

7.5 Salt Marshes 433

7.6 Sea Grass Ecosystems 440

7.7 Mangrove Ecosystems 442

7.8 Estuaries 443

7.9 Network Analysis 456

7.10 Potential Applications of the Ascendency Concept 469

7.11 Emergy Analysis 471

Appendix 473

References 477

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© 2001 by CRC Press LLC

CONTENTS

1.1 Introduction 1

1.2 Environmental Gradients and Stresses on the Shore 2

1.3 Salient Features of the Shore Environment 2

1.3.1 Seawater 2

1.3.2 Tides 4

1.3.3 Tidal Range and Proportions 4

1.3.4 Submersion and Emersion 7

1.3.5 Modifying Factors 9

1.3.5.1 Wave Action 10

1.3.5.2 Topography and Aspect 13

1.3.5.3 Climatic Factors 16

1.4 Patterns of Zonation on the Shore 16

1.1 INTRODUCTION

The relatively narrow strip where the land meets the sea provides the most diverse range of habitats for living organisms anywhere on the earth It offers a unique blend

of habitats not found elsewhere Animals and plants living there experience marked daily changes in environmental conditions as the tides ebb and flow These variations create stresses to which the species have adapted by organic evolution over long periods of time

The seashore is best defined as that part of the coast-line extending from the lowest level uncovered by the spring tides to the highest point washed or splashed by the waves This intertidal or littoral zone does not, how-ever, merge abruptly with the land above or the perma-nently submerged sublittoral zone below Above the high-est point reached by the waves, salt spray influences a zone which, on very exposed coasts, may extend several hundred meters up the shore Below the lowest point reached by the tides, a gradient in the quantity and quality

of light is reflected in the depth distribution of the plants and animals found there Furthermore, some sublittoral species extend up into the intertidal zone proper In this book, both of the transitional zones between the intertidal zone and the land above and the sublittoral below will

be considered

The extent of the intertidal zone depends on a variety

of factors, the most important of which are the angle of

slope of the shore and the amplitude of the tides This may vary from less than a meter on vertical rock faces where the tidal amplitude is small to several kilometers on soft shores where the tidal amplitude is high The physical features of this zone are of immense complexity, varying not only geographically, but from coast to coast in the same region and within comparatively short distances on any one shore Whether narrow or wide, this region is astonishingly rich in both the number of species of plants and animals and the densities of the individual species In the following pages we shall explore the complex shore environment, the ways in which representative species are adapted to life in a constantly changing environment, how they interact with each other, how the structure of the various communities is controlled, and how energy and materials cycle in the different ecosystems

Three interlocking factors determine the type of com-munity on the seashore: (1) the amount and intensity of wave action; (2) the type of substrate (whether rock, sand, mud, or some combination of these); and (3) the amplitude

of the tides To consider each of these in turn:

1 Waves are caused by wind, and their size is primarily determined by the uninterrupted dis-tance or “fetch” over which the wind can blow, the velocity and direction of the wind, and to a lesser extent the depth of the water Thus, the severity of wave action in any given locality is

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2 The Ecology of Seashores

determined by its geographical position in tion to the factors listed above As a conse-quence, there is every gradation in wave shock,from the pounding surf of exposed shores onthe western coasts of the main land masses tothe quiet waters of deep inlets, narrow estuaries,and fjords

rela-2 From a biological point of view, there are twomajor types of shores, “hard” and “soft,” theformer often referred to as rocky shores and thelatter as sand and mud shores The environmen-tal features and modes of life differ so muchbetween these two types that little overlap, ifany, exists between the species populationsinhabiting them There is, however, a gradationfrom rocky shores through boulder beaches,pebble beaches to sand beaches, and the lattergrade through muddy sand to mudflats Never-theless, it is convenient to preserve the distinc-tion between “hard” (rock and boulders) and

“soft” (sand and mud) shores

3 The third important factor, tidal exposure, will

be considered in detail later It is sufficient here

to note that the tidal range, or vertical distance,between high and low water not only varieswith the lunar cycle, but also geographically

The ebb and flow of the semidiurnal rise andfall of the tide result in periodic emersion andsubmersion of the intertidal zone, the extent ofwhich depends on a variety of factors (see Sec-tion 1.3)

1.2 ENVIRONMENTAL GRADIENTS AND STRESSES ON THE SHORE

Seashores are characterized by three main environmentalgradients:

1 The vertical (intertidal) gradient from sea to land

2 The horizontal gradient of exposure to waveaction

3 The particle size gradient from solid rockthrough boulders, pebbles, and coarse and finesands, to silts and clays

Across the intertidal gradient, there is a change fromthe highly stable, buffered subtidal environment to anunstable and increasingly stressed environment withincreasing tidal height Upshore the relatively stable envi-ronment changes until at the top of the intertidal zone(including the area influenced by wave splash and spray)

it merges with the unstable, terrestrial environment withdiurnal and seasonal changes in temperature, light, humid-ity, wind, and variable salinity due to rainfall and evapo-

ration Thus, across the intertidal gradient shore, isms are subjected to increasing stress

organ-As wave action increases along the horizontal exposuregradient, the upper shore is increasingly subject to wetting

by surging waves and spray, more food is transported tofilter-feeders, and better supplies of oxygen and nutrientsare available to the plants and animals On the other hand,increased stresses may be imposed on some organisms,e.g., problems of settlement, adhesion, and propensity fordislodgement and abrasive action of sand

The type of substrate, as discussed in Chapters 2 and

3, determines the type of community that can survive.These gradients interact in very complex ways withthe biological communities living on and in the sub-strate In the succeeding chapters we shall explore theseinteractions

1.3 SALIENT FEATURES OF THE SHORE ENVIRONMENT

When intertidal organisms are covered by the tides, theyare subjected to the same physiological conditions of tem-perature and salinity as those permanently submerged inthe sublittoral below Seawater from a physiological point

of view is a very complex solution The aspects of thiscomplexity that are of biological significance are ion con-centration, density, and osmotic pressure The osmoticproperties of seawater result from the total amount ofdissolved salts Seawater is composed of a number ofdifferent compounds that can be divided into the followingphases (Millero and Sohn, 1991):

1 Solids (material that does not pass through a0.45 µm filter)

a Particulate organic matter (plant and animaldetritus)

b Particulate inorganic material (minerals)

The Environment 3

can be conveniently grouped into major and minor stituents The major constituents, which are found every-where in the ocean in virtually the same relative propor-tions, are termed conservative elements The minorconstituents, on the other hand, show marked variations

con-in their relative concentrations due to selective removalfrom the water by living organisms, and are termed non- conservative elements.

The major constituents, comprising 99.9% of all solved salts, are sodium, magnesium, calcium, potassium,and strontium cations and the chloride, sulfate, carbonate,bicarbonate, and bromide anions, together with boric acidmostly in the undissociated state (Table 1.1) The mostimportant nonconservative constituents are the major plantnutrients, phosphates and nitrates, together with silicon,which is required by diatoms for the construction of theirfrustules and by radiolarians for their skeletons

dis-In addition to the dissolved inorganic substances, water, especially at inshore locations, contains appreciableamounts of organic material, both particulate (detritus)and dissolved The role of organic matter in materialcycling and in shore food webs will be dealt with in latersections of this book

The concentrations of the dissolved substances in water provide a means of determining the salt content ofseawater The total amount of inorganic material dissolved

sea-in seawater expressed as weight sea-in grams per kilogram (orparts per thousand) is termed the salinity, and is usuallyaround 35 gram kg–1, or 35 parts per mille Until the early1980s, salinity values were expressed in parts per thou-sand, or per mille, for which the symbol is “‰,” with theaverage salinity being 35‰ However, it is now standardpractice to dispense with the symbol “‰,” because asdetailed below, salinity is now defined in terms of ratio

The international standard method of estimating salinitywas based on determination by titration of the chlorinecontent (chlorinity) of the water (see Strickland and Par-sons, 1972), and the salinity was calculated from the rela-tionship: S‰ = 1.8065 Cl‰ (Sharp and Cuthbertson,1982) This definition assumes, among other things, thatall organic matter is oxidized, the carbonates are converted

to oxides, and the bromide and iodide have been replaced

by chloride Titration is time consuming, and various types

of conductivity salinometers are now used for the mination of salinity Thus, the practical salinity of a sam-ple of seawater is defined in terms of the conductivityratio, K 15 , which is defined by:

deter-at 15°C and 1 deter-atmosphere pressure, the concentrdeter-ation ofthe standard KCl being 32.3456 gram kg–1 The practicalsalinity is related to the ratio K15by a complicated equation.Other methods of determining salinity include therefractometer, which measures light refraction and thehydrometer, which measures density

In the open sea salinities commonly range from 32.00

to 37.00 The differences reflect local effects of evaporation,rain, freezing and melting of ice, or the influx of river water.Certain seas have markedly higher or lower salinities: theMediterranean, due to high evaporation and little freshwaterinflux, has salinities ranging from 38.40 to 39.00; the Baltic,which is a large brackish water body, ranges from 10.00 atthe mouth to 3.00 at the northern extremity

Regions where freshwaters mix with seawater aretermed estuarine In such regions, salinities undergo greatvariations depending on the state of the tide, the amount

TABLE 1.1 Concentrations of the Principal Ions in Seawater in Moles per Kilogram, in Parts per Thousand by Weight, and in Percent

of Total Salts Ion Moles kg –1

Percent by Weight

Percent of Total Salts

Chloride (Cl – ) 0.549 18.980 55.2 Sulfate (SO 42–) 0.0762 2.649 7.71 Bicarbonate (HCO3) 0.0023 0.140 0.35 negative ions Bromide (Br – ) 0.008 0.065 0.19 (anions) = 21.861%

Borate (H2BO3) 0.0004 0.026 0.07 Fluoride (F – ) 0.000004 0.001 0.0001 Sodium (Na + ) 0.468 10.556 30.40 Magnesium (Mg ++ ) 0.532 3.70 3.70 Calcium (Ca ++ ) 0.0103 0.400 1.16 positive ions Potassium (K ++ ) 0.0099 0.400 1.10 (cations) = 12.621%

Strontium (Sr ++ ) 0.0002 0.013 0.035 Overall total salinity = 34.482%

K15 Conductivity of the seawater sample

Conductivity of standard KCl solution -

=

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