Table Of ContentThe Geological Society of America
Memoir 67
TREATISE ON
MARINE ECOLOGY AND PALEOECOLOGY
Volume 1
ECOLOGY
Joel W. Iledgpeth, Editor
University of California, Scripps Institution of Oceanography
La Jolla, Calif.
Prepared under the direction of a Committee of the Division of Earth Sciences
National Research Council, National Academy of Sciences
Washington, D. C.
December 30, 1957
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COMMITTEE ON MARINE ECOLOGY AND PALEOECOLOGY
Harry S. Ladd, Chairman
Gordon Gunter, Vice-Chairman for Biology
Kenneth E. Lohman, Vice-Chairman for Geology
Roger Revelle, Vice-Chairman for Oceanography
Roland W. Brown Earl H. Myers
C. H. Edmondson A. W. B. Powell
K. O. Emery John B. Reeside, Jr.
Richard H. Fleming F. W. Rolshausen
Julia Gardner H. G. Schenck
Joel W. Hedgpeth Waldo L. Schmitt
Remington Kellogg John W. Wells
Frank C. Whitmore
Made in United States of America
Price $12.50
Address all communications to
The Geological Society of America
419 West 117 Street, New York 27, N. Y.
PRINTED BY WAVERLY PRESS
BALTIMORE, MARYLAND
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GEOL. SOC. AM., MEMOIR 67 (1) FRONTISPIECE
EDWARD FORBES (1815-1854) KARL MOBIUS (1825-1908)
From Wilson and Geikie: From a painting at Kiel:
Memoir of Edward Forbes, F. R. S. courtesy Adolf Remane
C. G. JOHANNES PETERSEN T. WAYLAND VAUGIIAN (1870-1952)
(1860-1928)
Photograph courtesy Gunnar Thorson
PIONEERS OF MARINE ECOLOGY AND PALEOECOLOGY
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The Memoir Series
of
The Geological Society of America
is made possible
through the bequest of
Richard Alexander Fullerton Penrose, Jr.
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Dedicated to
Thomas Wayland Vaughan
who organized the group responsible for this Treatise
and who served as a member until a few months before his death.
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Foreword
The late Dr. T. Wayland Vaughan, who had long been interested in marine ecology
and paleoecology, in 1940 planned and organized a Subcommittee on the Ecology of
Marine Organisms as a part of the Committee on Geologic Research (Norman L.
Bowen, Chairman) in the Division of Geology and Geography of the National Re-
search Council of the National Academy of Sciences. The members of this Subcom-
mittee were: C. H. Edmondson, Remington Kellogg, Harry S. Ladd (Chairman),
Kenneth E. Lohman, Roger Revelle, F. W. Rolshausen, H. C. Stetson, T. Wayland
Vaughan. The first annual report, which attempted to summarize current activities
in marine ecology and paleoecology, was issued in 1941. In this report the possibility
of preparing an ecological treatise was mentioned. In 1942 the Subcommittee was
made a full Committee and its title was changed to Committee on Marine Ecology
As Related to Paleontology to emphasize the paleontological nature of its interests.
With the encouragement and assistance of Walter H. Bucher, Chairman of the Divi-
sion of Geology and Geography, the Committee in its second report, issued at the
close of 1942, briefly listed its aims. There were six of these, the last of which read:
"Possibly, at a later date, to prepare a special treatise on ecology that would stress
geological interpretation." World War II greatly curtailed most activities in marine
ecology and also curtailed the activities of the Committee. Serious consideration was
given to the possibility of recessing the work, but the members felt that it might be
difficult to revive the program if work was stopped. Interest in the Treatise continued
at a low level until the war was over, but in 1946 in its sixth annual report the Com-
mittee published a tentative outline for the Treatise and invited criticism. In this
same year Kenneth E. Lohman was made Vice-Chairman of the Committee.
During the first 5 years of its existence the group that formed the original Sub-
committee continued to serve. Gordon Gunter was added as a member in 1942 and
Earl H. Myers in 1944. In 1946, the sixth year, H. C. Stetson resigned because of the
pressure of other duties, and three new members were added: K. O. Emery, of Cali-
fornia, A. W. B. Powell, of New Zealand, and H. G. Schenck, then stationed in Japan.
In 1947 four more members were added: R. H. Fleming, Julia Gardner, J. B. Reeside,
Jr., and Waldo L. Schmitt. At a two-day meeting held in March 1947, Gordon
Gunter was named Vice-Chairman for Biology, Kenneth E. Lohman Vice-Chairman
for Geology, and Roger Revelle Vice-Chairman for Oceanography; and concrete
plans for the completion of the Treatise were formulated. In 1948 Frank C. Whitmore
was made a member, and in the following year Roland W. Brown and Joel W. Hedg-
peth were added and assumed editorial duties. These additions brought the total
membership of the Committee to 19. In 1951 Doctor Vaughan asked that his name
be dropped from the Committee because of his failing eyesight. The Committee
accepted his resignation reluctantly, and John W. Wells was appointed to fill the
vacancy.
Most of the members of the Committee have prepared one or more units of the
text or the accompanying bibliographies, but the Committee alone could not have
produced the Treatise in its present form. To obtain as nearly complete coverage as
V
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vi FOREWORD
possible the aid of nearly 100 specialists was enlisted, each of whom prepared one
or more units in his particular field. To this large group the Committee extends its
deepest thanks. Formal titles of the contributors are not given but the field of interest
of each is indicated, together with his address at the time of going to press.
The Committee wishes to express its thanks to the Office of Naval Research, whose
support made it possible for Vice-Chairman Gunter to make an extended stay at the
University of California, Scripps Institution of Oceanography and a short visit to
the Woods Hole Oceanographic Institution in 1948-1949. Later, through a similar
co-operative arrangement between the Scripps Institution of Oceanography and the
Office of Naval Research, Joel W. Hedgpeth was able to spend several years at Scripps
assembling and organizing the materials for volume I and writing certain of the
units. During the summer of 1953, again with the support of the Scripps Institution
and the Office of Naval Research, he was able to travel in Europe and consult with
many of the contributors to the Treatise. The Committee also desires to express its
appreciation to the United States Geological Survey for encouragement and support;
six members of the Committee and many of the other contributors to the Treatise
are on the staff of the Geological Survey. Many members of the staff of the Geo-
logical Survey, the Scripps Institution, the National Museum and of other organiza-
tions have assisted the Committee in the critical review of manuscripts; this help is
gratefully acknowledged. We also wish to thank Karl P. Schmidt for translating
Chapter 25, which was submitted in German.
Messrs. Walter H. Bucher, W. W. Rubey, Arthur Bevan, Ernst Cloos, and Francis
Birch, who have successively served as Chairman of the Division of Geology and
Geography—now known as the Division of Earth Sciences—since the formation of
the Committee have encouraged and supported the work in every way possible. The
Committee desires to express its special thanks to Miss Margaret L. Johnson, Secre-
tary to the Division, for great assistance particularly in assembling and duplicating
of the Annual Reports, and to Mr. G. D. Meid, Business Manager of the National
Academy of Sciences, National Research Council, for aid in making publication ar-
rangements for the Treatise.
From the start the Treatise has been planned as an appraisal of accomplishments
in the fields of marine ecology and paleoecology, particularly those ecological in-
vestigations related directly or indirectly to paleontology. In attempting to obtain
broad coverage the Committee and its collaborators discovered several blank areas
in the paleoecological field and thus initiated several investigations that otherwise
might not have been started until some time later. These developments were gratify-
ing and have served to bridge the gap between past and present work in the field.
Paleoecology, though it deals with the past, is a comparatively youthful but active
branch of paleontology.
With these thoughts in mind it was decided to supplement the appraisals of past
work with some sort of prophetic look into the future. In what direction is paleo-
ecology going, how far may it hope to go, and what sorts of tools may be used in the
future? Some of the new techniques developed in recent years in borderline fields
such as biochemistry, biophysics, and geochemistry exhibit great promise and may
be successfully applied to paleoecological investigations. These prospects are dis-
cussed in a concluding chapter in Volume 2.
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CONTENTS
Page
Chapter 1. Introduction. By Joel W. Hedgpeth 1
Chapter 2. Classification of Marine Environments. By Joel W. Hedgpeth 17
Chapter 3. Concepts of Marine Ecology. By Joel W. Hedgpeth 29
Chapter 4. Obtaining Ecological Data in the Sea. By Joel W. Hedgpeth 53
Chapter 5. General Features of the Oceans. By Richard H. Fleming 87
Chapter 6. Solar Radiation, Submarine Daylight, and Photosynthesis. By Robert W. Holmes 109
Chapter 7. Salinity. By A. S. Pearse and Gordon Gunter 129
Chapter 8. Temperature. By Gordon Gunter 159
Chapter 9. Oxygen in the Ocean. By Francis A. Richards 185
Chapter 10. Carbonates and Carbon Dioxide. By Roger Revelle and Rhodes Fairbridge 239
Chapter 11. Nutrient Elements. By H. Barnes 297
Chapter 12. Biogeochemistry of Trace Metals. By Edward D. Goldberg 345
Chapter 13. Marine Biogeography. By Joel W. Hedgpeth 359
Chapter 14. Particulate Organic Detritus. By Denis L. Fox 383
Chapter 15. Interrelations of Organisms
A. Commensalism. By R. Phillips Dales 391
B. Parasitism. By Sewell H. Hopkins 413
C. Symbiosis. By C. M. Yonge 429
Chapter 16. Plankton. By Martin W. Johnson 443
Chapter 17. Bottom Communities. By Gunnar Thorson 461
Chapter 18. Rocky Intertidal Surfaces. By Maxwell S. Doty 535
Chapter 19. Sandy Beaches. By Joel W. Hedgpeth 587
Chapter 20. Coral Reefs. By John W. Wells 609
Chapter 21. Submarine Canyons. By Conrad Limbaugh and Francis P. Shepard 633
Chapter 22. Deep Sea and Abyssal Depths. By A. F. Bruun 641
Chapter 23. Estuaries and Lagoons. By K. O. Emery, R. E. Stevenson, and Joel W. Hedgpeth. 673
Chapter 24. Baltic Sea. By S. G. Segerstrale 751
Chapter 25. Black Sea and the Sea of Azov. By Hubert Caspers 803
Chapter 26. Caspian and Aral Seas. By L. A. Zenkevich 891
Chapter 27. Lunar Periodicity. By P. Korringa 917
Chapter 28. Fluctuations in Littoral Populations. By Wesley R. Coe 935
Chapter 29. Mass Mortality in the Sea. By Margaretha Brongersma-Sanders 941
ANNOTATED BIBLIOGRAPHIES
General Introduction. By Joel W. Hedgpeth 1011
Marine Borers. By Robert J. Menzies 1029
Marine Bacteria. By Claude E. ZoBell 1035
Marine Algae. By Maxwell S. Doty 1041
Nonalgal Marine Plants. By Edwin T. Moul and Roland W. Brown 1051
Marine Diatoms. By Kenneth E. Lohman 1059
Protozoa
Radiolaria. By William R. Riedel and Esther Aberdeen Holm 1069
Silicoflagellata. By G. Dallas Hanna 1073
Foraminifera. By Earl H. Myers and W. Storrs Cole 1075
Marine Sponges. By M. W. de Laubenfels 1083
Corals. By John W. Wells 1087
Recent Octocorals. By Frederick M. Bayer 1105
Recent Bryozoa. By Raymond C. Osborn 1109
Brachiopods. By G. A. Cooper 1113
Marine Worms. By Olga Hartman 1117
Oysters. By S. H. Hopkins 1129
vii
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viii CONTENTS
Nautilus. By H. B. Stenzel 1135
Mollusca (Except Oysters and Nautilus). By Willis G. Hewatt and Joel W. Hedgpeth 1143
Marine Crustacea (Except Ostracods and Copepods). By Waldo L. Schmit 1151
Marine Ostracoda. By Willis L. Tressler 1161
Copepods. By M. W. Johnson 1165
Xiphosura. By Carl N. Shuster, Jr 1171
Miscellaneous Arthropods. By Joel W. Hedgpeth 1175
Marine Insects. By Robert L. Usinger 1177
Crinoids. By Ailsa M. Clark 1183
Asteroids and Ophiuroids. By Elizabeth Deichmann 1187
Echinoids. By C. Wythe Cooke 1191
Holothurians. By Elisabeth Deichmann 1193
Chordates
Ascidians. By D. P. Abbott 1197
Pelagic Tunicates. By L. D. Berner 1201
Marine Cyclostomes. By Edward C. Raney 1203
Marine Fishes (Other than Cyclostomes). By Gordon Gunter 1205
Amphibians. By Karl P. Schmidt 1211
Reptiles (Except Turtles). By Karl P. Schmidt 1213
Sea Turtles. By Rainer Zangerl 1217
Birds. By Herbert Friedmann 1219
Marine Mammals. By Remington Kellogg and Frank C. Whitmore, Jr 1223
INDEX 1227
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Geol. Soc. America Memoir 67, Vol. 1, 1957
p. 1-16, 2 figs. Made in U. S. A.
Chapter 1
Introduction*
JOEL W. HEDGPETH
Biologist, University of California, Scripfs Institution of Oceanography, La Jolla, Calif.**
Ecology has no aim, but ecologists have. The problems of the ecologist are not fundamentally
different from those of any other kind of naturalist. The superficial differences in aim are due to the
different points of view, or methods of approach, rather than to any essential difference in the char-
acter of the problems.
—Charles C. Adams, Guide to the Study of Animal Ecology, 1913
Since the early history of ecology has been discussed in some detail by Allee el al.
(1949), among others, there is no need to repeat it here. Our concern is with the de-
velopment of marine ecology in general, especially as it may be related to paleo-
ecology. Gislen (1930) traced the study of "marine sociology," in particular of inter-
tidal zonation, as far back as 1812, although Wahlenberg's pioneer effort to describe
zonation was overlooked until 1917. The early work of d'Orbigny (1820) was also
for the most part neglected, and it is not until the decades 1830-1840 that we find
the unmistakable beginnings of modern marine ecology. In France it was the work
of Audouin and Milne Edwards (1832), whose scheme of zonation of the life of the
shore and shallow sea persisted for 50 years. In England, it was the influence of the
Manxman Edward Forbes that gave marine ecology its start. In Norway Michael
Sars, in Sweden Sven Loven, and in Germany Johannes Miiller turned their attention
to the animals of the sea. Dredging for bottom animals was the fashion of the day,
and in 1839 the British Association appropriated £60 to defray the expenses of "re-
searches with the dredge, with a view to the investigation of the marine zoology of
Great Britain, the illustration of the geographical distribution of marine animals,
and the more accurate determination of the fossils of the pleistocene period." Forbes
was the leading spirit on this dredging committee, and gave its activities a strong
ecological cast.
In Ireland an army surgeon, J. Vaughan Thompson, studied a number of strange
creatures that had usually been considered adult animals; he demonstrated that
they were really larval stages. These included the zoea of crabs and the nauplius
and cypris of barnacles. While Johannes Miiller is usually credited with "inventing"
the plankton net in the 1840's, Thompson was using such a net as early as 1828
(Hardy, 1953). Although Thompson published very little, his work was of the highest
quality : "It has been said of him that 'no great naturalist has written so little and
that so good.' " (Singer, 1950).
While many of the investigations of this period were concerned with zoology -per se,
* Contribution from the University of California, Scripps Institution Of Oceanography, New Series, No. 916a.
** Present address: Pacific Marine Station, College of the Pacific, Dillon Beach, Marin County, Calif.
1
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